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LECTURES
THE COMPARATIVE PATHOLOGY
OF INFLAMMATION.
KIP
LECTURES
ON THE
COMPARATIVE PATHOLOGY
OF INFLAMMATION
DELIVERED AT THE PASTEUR INSTITUTE
IN 1891
BY
METCHNIKOFF
/. ;
Chefde Service a C Institut Pasteur
TRANSLATED FROM THE FRENCH BY
F. A. STARLING AND E. H. STARLING, M.D.
WITH 65 FIGURES IN THE TEXT AND 3 COLOURED PLATES
LONDON
KEGAN PAUL, TRENCH, TRUBNER & CO., LTD.
1893
PREFACE.
THE outline of the natural history of inflammation here
offered to the reader, is not intended as an exhaustive
work on the pathology of inflammatory processes. My
principal object in writing this book being to show the
intimate connection that exists between pathology and
biology properly so called, I have purposely omitted
several points such, for instance, as the etiology of
suppuration, which has recently been so largely investi-
gated.
As the comparative anatomy of former times treated
only of man and the higher animals, so medicine has
hitherto excluded all the pathological phenomena
which occur in the lower animals. And yet the study
of these animals, affording as they do infinitely simpler
and more primitive conditions than those in man and
vertebrata, really furnishes the key to the comprehen-
sion of the complex pathological phenomena which
are of special interest in medical science.
If we examine the processes of inflammation from
this point of view, we shall be able to form a more
complete and definite idea of their real significance.
In the following treatise, this theoretical side is indeed
the only one considered ; but here again, as in so many
VI PREFACE.
other cases, the extension of theoretical knowledge must
inevitably react upon its practical application.
The subject of phagocytes is frequently referred to
during the course of this study on the biological theory
of inflammation. I must however warn the reader
that he will not find a full exposition of the phagocyte
theory in this work. Many points in connection with
this question come under the headings of immunity,
regeneration and atrophy points which I hope to treat
separately later on.
With the exception of some slight modifications,
these lectures are published in the form in which they
were delivered at the Pasteur Institute, in April and
May of 1891. I must, however, mention that in deal-
ing with the destruction of the tubercle bacillus in the
organism (Lecture X.), I have made use of the reactive
phenomena presented by the tuberculous cells of
Meriones instead of describing those in Spermophilus,
as I did in my course. Rather than repeat here what
has already been published in a separate paper, I have
preferred to introduce an example which has not yet
appeared in print.
In conclusion, I must express my great indebtedness to
Professor H. de Lacaze-Duthiers, who has supplied me
with valuable material for the study of inflammation
in the lower animals, and also to my colleagues at
the Pasteur Institute, Professor Duclaux and Dr. E.
Roux, who have spared themselves no trouble in help-
ing me in the preparation of this work.
ELIAS METCHNIKOFF.
Paris, January 15, 1892.
PREFACE TO THE ENGLISH
EDITION.
I WAS much gratified when the proposal was made to
me of publishing an English edition of this study on
inflammation, the more so as in it I have endeavoured to
apply to pathology the principles of evolution which we
owe to the genius of English philosophers. I have indeed
dared to put forward anew theory of inflammation, only
because I felt that I had Darwin's great conception as a
solid foundation to build upon, supported as it is by the
doctrine of natural selection. The biological theory of
inflammation has already endured the test of time. It
was first advanced in several papers, the earlier of
which were published about ten years ago and have been
the subject of manifold attacks and criticisms. The
appearance of the French edition of this work, about a
year ago, was the signal for a renewal of the objections
to the theory, especially by German workers. These
criticisms I have endeavoured to meet in an article
printed as appendix to this volume, and have shown
that all the objections which have been brought for-
ward bear only on points of subsidiary importance,
without touching the root of the question, and indeed
in many cases rest on simple misunderstandings which
have arisen through the inadequacy of my treatment.
viii PREFACE.
In replying to my critics, I have not mentioned some
objections which have been raised by English patho-
logists. The most important feature in these is the
idea that it is principally the eosinophile cells which
emigrate through the vessel-wall in inflammation ; that
is to say, a variety of leucocytes which never acts as
phagocytes. I have already replied to this statement in
a special article (Annales de PInstitut Pasteur, No. I,
1893), and need not repeat all my arguments here. I
would only observe that the leucocytes which escape from
the vessel in inflammation consist for the greater part of
true phagocytes. Special investigations, which my
pupils and I have carried out in my laboratory
on the eosinophile cells, have shown conclusively that
under no circumstances have the eosinophile granules
the microbicidal action which has been attributed to
them by some pathologists. These granules rather re-
present reserve materials, exactly similar to yolk granules
and aleurone grains, which are also eosinophile.
I would conclude by expressing the hope that the
history of the evolution of inflammation which is here
put forward may be found to withstand the test of any
further criticisms that may be directed against it ; and
I trust that the book in its present version may be as
favourably received as has been the pleasant fate of the
original.
ELIAS METCHNIKOFF.
Paris, April loth, 1893.
CONTENTS.
PAGE
PREFACE v
PREFACE TO THE ENGLISH EDITION vii
LECTURE I.
Infection a struggle between two organisms Instance of the
Sphaerophrya ComparativePathology abranch of Zoology
Elementary principles of this science Inflammation
General survey of the principal theories in relation to this
phenomenon Present views on the subject Necessity of
adopting the comparative method in the study of inflamma-
tion i
LECTURE II.
Are the unicellular organisms subject to traumatic and in-
fectious maladies ? Merotomy of the Amoebae and Infusoria
Lesions of Vaucheria Epidemic disease of Amoebae,
caused by the Microsphaara I ntracellular digestion in the
Protozoa Digestion of bacteria Epidemics in the Infu-
soria : disease of the nucleus and nucleolus Division of
infected Paramaecia, and the means whereby they rid
themselves of the parasite Acinetas Chytridia . .14
LECTURE III.
Plasmodium of Myxomycetes Puncture by a glass tube
Cauterisation Chemical excitation Trophotropism
Chemiotaxis Habituation of the plasmodium Negative
chemiotaxis Repulsion of the plasmodium in the presence
of bacteria Digestion of bacteria by the plasmodium
Sensibility of the plasmodium Fixed plants having no
a
X CONTENTS.
PAGE
true intracellular digestion Necrosis and regeneration
Waldenburg's experiments Functions of the cell-wall
De Bary's observations on Peziza sclerotiorum Tumours
of plants . 29
LECTURE IV.
Transition from the unicellular organisms to the Metazoa
Sketch of the phagocytella theory Protospongia
Sponges : their organisation their three layers their
nutrition Intracellular digestion Ablation of parts of
the Sponge Artificial division Introduction of pointed
bodies Utilisation of foreign bodies to assist in forming
the skeleton Fate of the organisms which have pene-
trated into the interior of the Sponges Protective function
of the ectoderm Comparison with the Myxomycetes
Comparison with the inflammation of vertebrata . . 42
LECTURE V.
Cwlenterata, Echinodermata and Verities Traumatism and
regeneration in Hydra Accumulation of phagocytes in
Acalepha (Scyphomedusae) Phagocytes of star-fishes
Inflammation in Bipinnaria Reactive changes in the
perivisceral cells of the Annelida Phagocytic reaction in
the diseases of Nais and Lumbricus Struggle between
the phagocytes of Lumbricus and Rhabditis Microbic
infections of Worms 56
LECTURE VI.
Arthropoda, Mollusca, and Tunicata Their vascular system
Their Phagocytes Spleen of the Gasteropoda In-
flammatory reaction Diapedesis in intact Ascidians
Introduction of bacteria into the body of Ascidians and
Crustaceans Infectious disease of sandhopper (Talitrus)
Diseases of Daphnia Introduction of bacteria into
insects Epidemics among insects 75
LECTURE VII.
Vertebrata Amphioxus Embryos of Axolotl Young larvae
of Urodela Comparison with the invertebrata Tadpoles
Diapedesis Migratory cells Fixed cells Phagocytic
properties of leucocytes Do fixed cells also functionate as
CONTENTS. xi
^f PAGE
phagocytes? Transformation of leucocytes into fixed
connective tissue cells Fate of the leucocytes that do not
undergo this transformation Evolution of inflammation
in the organic world 94
LECTURE VIII.
Varieties of leucocytes Origin of these varieties Mobility
^Phagocytic properties Condition of englobed microbes
Their vitality and virulence Sensibility of leucocytes
Tactile sensibility Chemiotaxis Buchner's investiga-
tions Leucocytosis Intracellular digestion Destruction
of microbes, especially in immune animals Action of
leucocytes on spores Multiplication of leucocytes by
direct and indirect cell-division Changes they undergo
Transformation of lobed into single nuclei . . .in
LECTURE IX.
Endothelium of vessel walls Their development out of the
mobile cells of the embryo Development of the capil-
laries Contractility of the endothelial cells Star cells
Phagocytosis in the endothelial cells Fixed connective
tissue cells Clasmatocytes of Ranvier Ehrlich's cells
Active migration of leucocytes in diapedesis Experi-
ments with quinine (Binz, Disselhorst) " Itio in partes ''
Dilatation of the vessels Theory invoking the influence of
the surrounding tissue Influence of the nervous system
Negative chemiotaxis of the leucocytes in cases of severe
infection 137
LECTURE X.
Chronic inflammations Tuberculosis as a type of a chronic
inflammation Phagocytic nature of tuberculous cells
Destruction of tubercle bacilli by phagocytes Power of
resistance of Merioties to tubercle virus Leprosy . . 157
LECTURE XI.
Serous inflammations Two classes of these inflammations
Bactericidal power of the humours and the serous exsuda-
tions Antitoxic property of serum and the serous in-
flammations 172
xii CONTENTS.
LECTURE XII.
PAGE
Review of other theories of inflammation in light of the ac-
quired facts Nutritional theory of Virchow Vascular
theory of Cohnheim Experiments of the latter on the
tongue of the frog. Introduction of irritative agents into the
blood. Argument against Cohnheim's theory furnished by
the reaction in vertebrata Struggle of the organism with
external agencies Use of intracellular digestion Phago-
cytes Haemitis (recurrent fever, disease of Daphnia)
Tuberculosis. Essential nature of inflammation Sensi-
bility of the phagocytes Its progressive development
Sensibility of the endothelial cells Definition of inflamma-
tion. Inflammation is not regeneration Inflammation is
not identical with resorption Objections raised to the
biological theory of inflammation Vitalism Teleology
Absence of phagocytes in certain infective lesions Im-
perfections in the inflammatory reaction Surgical inter-
ference Comparative pathology 180
DESCRIPTION OF THE PLATES 199
APPENDIX. A CONSIDERATION OF SOME CRITICISMS ON
THE BIOLOGICAL THEORY OF INFLAMMATION . .201
INDEX 213
THE COMPARATIVE PATHOLOGY OF
INFLAMMATION.
LECTURE I.
Infection a struggle between two organisms Instance of the Sphae-
rophrya Comparative Pathology a branch of Zoology Ele-
mentary principles of this science Inflammation General
survey of the principal theories in relation to this phenomenon
Present views on the subject Necessity of adopting the
comparative method in the study of inflammation.
IN deciding to give a few lectures on a subject belong-
ing to the domains of pathology, I have resolved to do
so solely in my capacity of zoologist. The complexity
of the most important pathological processes, as studied
according to the universal custom on vertebrates, is so
great, even in so low a member as the frog, that it
becomes impossible to analyse them or to attain any
adequate conception of their real significance.
It is unnecessary to cite any special proof in support
of the doctrine that disease and pathological processes
are evolved in the same way as man and the higher
animals themselves. In all organisms, starting from the
simplest forms of life, we find infectious diseases pro-
duced by different classes of parasites. It is therefore
only natural to suppose that this parasitism gives rise
B
INFLAMMATION.
to a definite series of disturbances in the infected organ-
ism, and likewise provokes phenomena of reaction in
the latter.
If we examine the organisation of an animal or a
plant, we find that their most characteristic features are
their organs of attack and defence. The carapace of
the crayfish, the shell of molluscs and the teeth of the
vertebrates, as well as many other organs, are so many
means of protection to these animals in their perpetual
warfare. The mere enumeration of all the organs
acquired for the
purpose of helping
them in this strug-
gle would involve a
complete account
of the comparative
anatomy of ani-
mals.
Now from active
aggression to in-
fection, there is but
a short step. To take an instance from the lower
animals, we have only to consider the biological rela-
tions between some kinds of Infusoria. Among these
minute animals there is a group of suckers, which put
out their feelers in all directions, with the object of
attacking other varieties of Infusoria and of absorbing
their contents. (Fig. I.) Most of these Acinetae are
Infusoria attached to different objects which occur in
water, and are entirely predatory in their habits.
Observation of their behaviour will explain many points
in their organisation, and will also throw light on the
mode of resistance offered by the infusoria they attack.
Fig. i. Spheerophrya niagna (after Maupas).
LECTURE I.
Among these Acinetae there are some which may be
distinguished by their excessive minuteness, as well as
by the fact that, instead of being attached to some
object and drawing their prey towards them, they are
free and fasten themselves on to other infusoria larger
than themselves. These minute
Acinetae pierce the outer covering
of the Infusoria they attack, and
take up their abode in the proto-
plasm of their host where they
lead a parasitic existence. (Fig. 2.)
We see then that organisms
which are very nearly related to
each other in fact two members
of the same class (Sphceroplirya
magna and Sphcerophrya para m<z-
cioruni) may act as voracious
aggressors or as parasites with the
power of producing a definite in-
fection. Since zoological research
takes cognisance of the phenomena
of attack and defence, it should
likewise include the processes of
infection and resistance, which are
really in such close connection
with the former. The phenomena
of the active struggle among animals, however, being
much more prominent, havejattracted the attention of
naturalists for years, whereas those of infection, which
are far less on the surface, have been but rarely and in-
sufficiently studied.
A branch of zoology should be devoted to the in-
vestigation of the various powers of adaptation by
B 2
Fig. 2. Sphcerophrya. para-
a. Two parasitic sphaero-
phryae.
b. A parasite adherent to
the surface of the host.
4 INFLAMMATION
means of which an animal may penetrate and remain
in the body of other animals, as well as to the special
study of the reactive phenomena and mechanisms
which serve organisms in their resistance to parasitic
invasion. A branch of zoology the comparative
pathology of animals may thus be formed, which
would differ from the present comparative pathology in
many ways. Whereas the latter, which has been mainly
founded on veterinary science, is only concerned with
the higher animals and chiefly with mammals, genuine
comparative pathology should include the whole animal
world, and be treated from the widest biological stand-
point.
The groundwork of such a comparative pathology
was laid about five-and-thirty years ago. About this
time, in 1857 and 1858, the theory of natural selection
was built up on scientific foundations by Darwin and
Wallace, the biological theory of fermentation by
Pasteur, and the theory of cellular pathology by
Virchow.
The first of these theories, which now forms the basis
of all biological research, proved the genealogical
evolution of organized beings, and explained the adap-
tation of means to an end observed in them. It de-
monstrated that only the characteristics which are
advantageous to the organism survive in the struggle for
existence, while those that are harmful to the individual
are readily eliminated by natural selection.
The biological doctrine of fermentation, established
by Pasteur with the discovery of the lactic ferment in
1857 an d with that of the butyric ferment in 1861,
showed at once in what direction the causes of infection
should be sought, so that the forgotten discovery of the
LECTURE I. 5
anthrax bacillus was readily brought to light again by
Davaine, and became the starting point of pathological
bacteriology.
When Virchow demonstrated the important part
played by the cells of the organism in pathological
processes, a third link was formed in this chain of bio-
logical theories, which are indispensable to the founding
of a true comparative pathology.
But although the groundwork of this science was laid
more than thirty years ago, we are not yet in a position
to treat questions of general pathology from a com-
parative standpoint. An inquiry into the pathological
doctrines which are at present held concerning the most
important morbid processes, will readily prove the truth
of this statement.
As an example we will take inflammation, which is
universally admitted to be the most important phe-
nomenon in pathology. We will first examine the
results obtained by the usual methods of investigation,
and then we will proceed to inquire whether it would
not be more desirable to treat the question of inflam-
mation according to the comparative method.
As it is impossible to give a complete account of all
the theories on the subject of inflammation, we will
briefly consider those which are the most generally
accepted and taught by pathologists.
For a considerable time it was principally the ap-
pearance of inflammation the ' rubor ' which at-
tracted the attention of medical men, who were thus
frequently induced to regard hyperaemia as the most
essential act in inflammation, and even to identify these
two conditions. The theories concerning inflammation
were thus reduced to an analysis of the hyperaemia,
O INFLAMMATION.
which was supposed to be caused either by paralysis of
the vasomotor nerves (paralytic theory) or by a spas-
modic contraction of the affected arteries, accompanied
by an afflux of the blood to the neighbouring parts
(spasmodic theory).
But it was soon seen that hyperaemia of itself is in-
capable of producing true inflammation. A temporary
hyperaemia, or even one of longer duration, may occur
without resulting in exsudation, which is an essential
condition in cases of typical inflammation. In order to
explain the ' tumor/ it was suggested that the tissues
at the seat of lesion had an attractive influence on the
blood. This view was more definitely formulated in
Virchow's conception of an increased nutritive and repro-
ductive activity of the cells at the seat of inflammation,
which gave rise to the formation of a large quantity of ex-
sudation cells at the expense of the cells of the damaged
tissues. According to this theory, hyperaemia would be
merely a subordinate and entirely secondary phenomenon.
The definite proof given by Cohnheim, that the cells
in inflammatory exsudations arise from the white cor-
puscles of the blood, first enabled pathologists to decide
accurately upon one of the principal questions in con-
nection with inflammation. After having established
this undoubted fact, Cohnheim adopted the opinion of
Samuel, according to whom the main factor in all in-
flammatory states consists in a lesion of the vessels
which are attacked by the irritating cause. The
inflamed vessels, being more permeable, allow the fluid
and corpuscular elements of the blood to flow through
them in a purely passive manner. These exsuded
products collect at the part where they meet with least
resistance, and thus produce the inflammatory tumour.
LECTURE I. 7
In this theory the tissues at the point of lesion, as
well as the hyperaemia and the vasomotor phenomena,
play a very unimportant part. Although certain ideas
in connection with this theory have met with more or
less serious objections, it is accepted, especially in
Germany, by the majority of contemporary pathologists
who have formulated any general conception of inflam-
matory processes. Some authorities adopt it in general
terms, while laying stress at the same time upon the
importance of the phenomena occurring in the damaged
tissues and the vasomotor system. As it is not possible
to bring these conditions into connection with one
another, they content themselves with a simple enume-
ration of the changes resulting from inflammation which
take place in the tissues and the vascular system.
Ziegler, 1 who is the author of the best known work on
pathological anatomy at the present time, admits in his
chapter " On the Definition of Inflammation/' that he is
unable to give a clear definition of this condition. In
his own words : " The idea of ' inflammation ' includes a
whole series of phenomena, which occur partly in the
circulation and partly in the tissues, and may be com-
bined in various ways. As we are not here dealing with a
simple pathological condition, it is impossible to give a
short concise definition of inflammation. Even if only
certain phenomena, such as those which occur in the
circulation, were taken as characteristic of inflammatory
processes, a definition of them would certainly fail to
convey an adequate conception of inflammation." Ac-
cordingly, Ziegler merely gives a summary account of
the changes produced by inflammation.
1 " Lehrbuch der patholog. Anatomic," 6th edition, 1889, vol. i.
p. 1 86.
8 INFLAMMATION.
Recklinghausen l holds that it is " at present impossible
to determine the primum movens, the starting point of
the changes ; that is to say, the site of the earliest
lesion." So he, also, is unable to do more than give
a detailed and careful description of inflammatory
phenomena.
The definition of inflammation put forward by Cornil
and Ranvier 2 consists simply in an enumeration of the
events in this pathological process. They define it as a
" series of phenomena, observed in the tissues or organs,
and analogous to those artificially produced in the same
parts by the action of a physical or chemical irritant"
In order to simplify this complicated question, it was
endeavoured to investigate this pathological condition
in parts of the body devoid of blood-vessels, in which
inflammation of the tissues only could be observed.
Attention was concentrated upon the cartilages, and
mesentery, and especially upon the cornea. From the
changes observed in the cells of these organs, it was
argued that vascular disturbances were not essential
for the production of inflammatory phenomena in the
tissues. These changes consist in proliferation of the
local cells, and their return to " the embryonic con-
dition."
Cohnheim, however, in his experiments on the cornea,
demonstrated vascular intervention in experimental
keratitis, and proved that immigration of leucocytes com-
ing sometimes from the margin of the cornea and
sometimes from the conjunctiva, took place into the
1 "Handbuch d. allgemeinen Pathologic des Kreislaufs," 1883,
p. 198.
2 " Manuel d'histologie pathologique," 2nd edition, vol. i. p. 94.
LECTURE I. 9
seat of inflammation. These results showed at the same
time that the attempts to eliminate vascular influence,
even in the most evascular organs of the higher animals,
had been quite futile.
An endeavour was then made to bring the changes
produced within the cells themselves into the cycle of
inflammatory phenomena, and Virchow's theory of par-
enchymatous inflammation was reinstated. The limits
of inflammation were thus considerably extended.
Brault l and others have gone even further, and sought
to include under this term the acute degenerative phe-
nomena of the cells.
We see, therefore, that the attention of pathologists
has of late been mainly concentrated upon the part
played by the vascular system and the local tissue-
elements respectively. By the discovery of the pheno-
mena of karyokinesis, which prepared the way for the
solution of many problems dependent on the formation
and origin of the cells, a new life has been given to
the discussion as to whether the inflammatory cells
originate at the expense of the white blood corpuscles,
or in consequence of the proliferation of local cells. A
discussion on this subject has recently been raised
by Grawitz, 2 an ardent disciple of Virchow's, who
maintains that a large proportion of pus-globules are
formed from the cells of the connective tissue, and by
Weigert, 3 a faithful pupil of Cohnheim, who upholds
the main theory of this pathologist with regard to the
origin of the inflammatory cells from the leucocytes
1 "Etude sur 1'Inflammation," Paris, 1888, p. 34.
2 Deutsch. med, Wochensch., 1889, No. 23.
3 Fortschritte der Median, 1889, Nos. 15 and 16.
IO INFLAMMATION.
that have escaped through the vessel walls. The dis-
covery of the karyokinetic phenomena enabled ob-
servers to ascertain, beyond a doubt, that very frequent
division of the local tissue-cells takes place at the
seat of inflammation. But whereas the partisans of
Virchow's doctrine regarded this as a proof of the part
played by these local elements in the formation of the
inflammatory tumour, the adherents of Cohnheim's
views interpreted this cell division as a simple pheno-
menon of reparation, serving to restore the mischief pro-
duced by the primary lesion. As this view became
more completely accepted, it gave rise to a distinction
of two classes of phenomena in inflammation ; first, in-
flammation properly so called, i.e. the lesion of the
vessel walls and other disturbances brought about by
the irritating cause ; secondly, reparation, consisting in
the regeneration of the missing tissues and in the
formation of the scar. The most advanced exponent of
this classification, Roser, 1 went so far as to assert that
inflammation is a true disease, due to the infection by
microbes, and that the reparatory phenomena con-
stitute its' cure. According to this authority it is even
impossible " to give a single definition of inflammation
so long as this name is held to include the most hetero-
geneous phenomena, such as the disorders due to infec-
tion and the processes of recovery."
But besides this mode of interpreting inflammation, a
movement has been for many years travelling in the
opposite direction. Instead of separating inflammatory
phenomena into two fundamentally distinct classes, they
have been regarded as stages of a single process repre-
1 "Entziindung und Heilung," Leipzig, 1886, pp. 9, 11, etc.
LECTURE I. II
senting a salutary reaction against some injurious influ-
ence. According to this doctrine, not only regeneration
and cicatrization but also the primary processes of
inflammation, such as emigration and the alteration in
the vessel walls, would be considered as reparatory acts
serving to counteract the damage brought about by the
irritant. This theory, which was clearly stated by L.
Sachs more than fifty years ago, has found fresh sup-
porters at different times. It was accepted by Buchner '
in general terms and it has recently been developed
by Neumann, 2 who maintains that true inflammation
never occurs except where there has been a primary
lesion of the tissues. The definition of inflammation
which he endeavours to introduce, is as follows.
" Under this name we should include the series of
local phenomena which are developed as a result of
primary lesions of the tissues (l<zsio continui or necrosis],
and tend to cure them." 3
From this review of the present state of our know-
ledge on the subject of inflammation, it is apparent that,
in spite of all the pains taken by investigators, the
methods hitherto adopted are inadequate for the study
of phenomena so intricate and variable in their mani-
festations. It is not to be wondered that several
authors, as, for instance, Thoma, 4 should have proposed
to suppress the term ' inflammation ' altogether.
In spite of many attempts to simplify the experimental
conditions and to eliminate certain factors from this
1 " Prophylactische Therapie der Lungentuberculose," 1882.
2 " Ueber den Entziindungsbegriff," Ziegler's Beitrage zurpatho-
logischen Anatomic, 1889, vol. v. p. 347.
3 Loc. tit., p. 363.
4 Berliner klinische Wochenschrift, 1886.
12 INFLAMMATION.
complex process, we have only succeeded in the case
of the elevation of temperature. By studying inflamma-
tion in cold-blooded animals incapable of generating
heat to any appreciable extent, such as frogs, it is seen
that true inflammation can take place in the total ab-
sence of heat, one of the four classical factors (dolor,
calor, rubor, tumor). In these animals the inflamma-
tory nature of the phenomena in question is so evident,
that no one has raised any objection to the application of
the term ' inflammation ' in cases where the temperature
is not raised and where, consequently, the word is no
longer applicable in its etymological sense.
The frog has always been chosen on account of the
facilities it presents for experimental study, without
realizing that in so doing a method of comparative
pathology was adopted. It is along these lines that
investigation should proceed, but the comparison
should be extended to still lower members of the
animal kingdom, in order to eliminate further factors
and to study these phenomena in a yet more simple
condition.
As we have already seen, all attempts to obtain
inflammation in the higher animals without the inter-
vention of the vessels have failed, since it is impossible
to exclude the circulatory system, even in the most
isolated tissues. In order to obtain definite results, we
must direct our attention to the large field presented by
the invertebrata, among which there are many animals
completely devoid of blood vessels. The comparative
method has already rendered good service, not only in
the realm of natural sciences, properly so called, but
even in the study of the most complicated phenomena.
Thus Psychology owes much to the observation of
LECTURE I. 13
psychical phenomena in the lowest animals, and even
in the social sciences, such as ethnology or political
economy, investigation has often been extended to
the most inferior races. Pathology is almost the only
science in which the comparative method has been
ignored, although it has to do with phenomena which
present complications from every point of view, and it
would be particularly profitable to make use of such
methods for enlarging the scope of its investigations.
The question may be thus formulated. Do the
factors (traumatic or infective) which evoke the series of
phenomena known as inflammation in man and the
higher animals, produce any analogous conditions in
the lower vertebrata, such as Amphioxus, or in the
invertebrata ? Is the existence of a circulatory system
essential for the setting up of inflammation, or does
this also occur in animals which possess no blood
vessels, and in this case how does the nervous system
act ? For inflammation to take place, is it necessary
that the animal should possess a certain number of
differentiated organs or may it consist merely of an
agglomeration of non-differentiated cells ? Do we find
anything analogous to inflammation in plants? Are
there any instances of inflammatory action in uni-
cellular organisms ? In the following lectures we shall
discuss these questions one by one, and endeavour to
answer them.
LECTURE II.
Are the unicellular organisms subject to traumatic and infectious
maladies ? Merotomy of the Amoebae and Infusoria Lesions
of Vaucheria Epidemic disease of Amoebae, caused by the
Microsphaera Intracellular digestion in the Protozoa Diges-
tion of bacteria Epidemics in the Infusoria : disease of the
nucleus and nucleolus Division of infected Paramaecia, and
the means whereby they rid themselves of the parasite
Acinetae Chytridia.
WE will first inquire whether the unicellular organ-
isms, which abound in the media surrounding us, are
subject to infectious diseases, and whether they are
susceptible to those influences which produce in us a
more or less pronounced inflammation. We shall after-
wards examine the changes that these influences call
forth in these lowly organisms.
In man and the higher animals a traumatic lesion,
even when insignificant, invariably provokes the series
of phenomena which characterize inflammation. In
unicellular organisms the resulting events are much
simpler. If we cut an Amoeba in two, there is not even
a wound formed along the line of section, for the edges
unite immediately after the passage of the instrument.
(Figs. 3, 4.) Two new Amoebae are thus produced ; the
one which encloses the nucleus continues to grow and
behaves in all respects like a normal individual, while
the other, which is without any nucleus, dies at the end
LECTURE II. 15
of a longer or shorter period. 1 Some other inferior
organisms, which contain several nuclei, as for in-
stance Actinophrys, can be divided into several
pieces, each of which is regenerated in a very short
time, provided that it still contain a fragment of
nucleus. 2 In the Infusoria, which possess a more
highly differentiated protoplasm, artificial bisection
Fig. 3. An amoeba immediately after
bisection.
a. The half containing the nucleus n.
b. Half without nucleus.
v. Contractile vesicle.
Fig. 4. The same amoeba five minutes
after the operation.
(After Bruno Hofer.)
produces a wound which lays bare the inner layer of pro-
toplasm. After a short time, however, the edges of the
peripheral layer grow over the wound, and secrete a new
cuticle, thus securing complete cicatrisation. (Fig 5.)
1 Bruno Hofer. " Experimentelle Untersuchungen iib. d. Ein-
fluss des Kerns auf das Protoplasma." Jenaische Zeitschr.
f. Naturwiss., 1889, vol. xxiv. p. 109, pi. iv. and v.
2 K. Brandt. " Ueb. Actinosphaerium Eichhornii," 1887, p. 30.
16
INFLAMMATION.
These phenomena are almost exactly the same
whether the fragments be provided with a nucleus or
not. In those with a nucleus, however, regeneration is
complete in a very short time (often in less than twenty-
four hours), while the others gradually atrophy and in
the end always die. Balbiani, 1 who has published an
important paper on the merotomy of the Infusoria, is
indeed of opinion that cicatrisation is never properly
completed in the fragments without a nucleus, the
latter exercising a decided influence on the secretion
Fig. 5. Merotomy ofa Stentor,
a. Anterior fragmemt ; m, middle fragment ; f, posterior fragment ; wz 1 , ; 2 , m 3 , m*,
stages in the regeneration of the middle fragment.
(After Balbiani.)
of the cuticle. In some species, such as Trachelius
ovum, the wounds caused by the section are imme-
diately covered over by the ectoplasm, and the separate
fragments that still possess a nucleus are completely
regenerated in less than five hours.
In the same way unicellular plants may undergo
severe injuries without necessarily perishing in conse-
quence. Thus Hanstein 2 has observed that, when a
1 " Recherches exp^rimentales sur la m^rotomie des infusoires
cilies." Recueil zoologique suisse, vol. v. 1888.
5 Vide Frank, " Die Krankheiten der Pflanzen," 1880, vol. i.
P- 97-
LECTURE II. 17
part of the unicellular alga Vaucheria is cut or
crushed, it is only the damaged part that dies, while
the rest of the cell is healed by the secretion of a
cuticular layer on the injured surface, and the forma-
tion of a sort of sequestrum.
In these phenomena then, in the lower organisms, we
have to do simply with a regeneration that takes place
more or less completely and readily. But after trau-
matic lesions it is in-
fection that most fre-
quently provokes in-
flammation. Now, in-
fectious diseases are
very common among
the Protozoa and uni-
cellular plants. Even
the lowest members of
these classes are some-
times subject to infec-
tion.
Thus in the Amoebae
I have observed an Fig. 6. Amoeba infected by the Microsplurra.
... , , a. Early stage.
epidemic caused by a
very simple organism, which occurs in the form of
a round cell provided with a very delicate wall and
a nucleus, and capable of multiplying by division.
The large Amoeba with rounded pseudopodia, that
feeds upon diatoms, sometimes contains by the side of
these brown algae a small number of these round cells
(Fig. 6), which I shall allude to under the name of
Microsphara. As the general aspect of the amoeba
together with its protoplasmic movements remain
normal, no one would suspect this rhizopod to
C
i8
INFLAMMATION.
be diseased. Continued observation shows, however,
that whereas the enclosed diatoms undergo digestive
changes, the Microsphaerae divide and multiply without
let or hindrance within the protoplasm of the amoeba.
This latter ejects the diatoms and becomes gradually
less and less active, showing that it is not in a healthy
condition. At the same time the protoplasm becomes
filled with Microsphaerae, and the Amceba invaded by
the parasite finally perishes. (Fig. 7.)
This case is in-
teresting since it
shows us that an
organism, although
composed almost
entirely of a proto-
plasm which has
the power of readi-
ly digesting the
contents of dia-
toms, can never-
Fig 7. A dying Amoeba, full of parasitic Microsphsera:. theleSS be infected
by another organ-
ism. The infecting agent, which is to all appearance
insignificant, has yet the power of resisting the digestive
influence of the Amceba and of bringing about its death.
To explain this fact, we must assume some property in
the parasite enabling it to produce in the interior of the
amoeba a substance which protects the Microsphasra,
and is at the same time toxic for the amoeba.
Thus the infection develops in spite of the marked
power of intracellular digestion possessed by the
Amoebae. A closer observation of the group of
Protozoa compels us to the conviction that this
LECTURE II. 19
digestive function must play an important part in the
mutual relations of these lowly organisms. Many
Rhizopoda and Infusoria live in media swarming with
other unicellular organisms, including bacteria. The
latter, which multiply very rapidly, serve as food to
many of the Protozoa. Thus various Amoebae devour
bacilli, which undergo certain definite changes in the
interior of their protoplasm. Without altering their
shape, the bacilli acquire the power of taking up solu-
tions of vesuvine, which does not stain these microbes
when living in their natural conditions. (Fig. 8.) Since
Fig. 8. An Amoeba living in the midst of bacilli of which
it has taken up a certain number.
precisely similar changes are also observed in the interior
of Vorticellae and other Infusoria which live on bacteria,
it is evident that they are due to a digestive influence
exerted by the contents of the Protozoa. This conclu-
sion is in harmony with the observation of B. Hofer * on
digestion in Amoebae. This investigator has shown that
the more the food is altered in the interior of these Rhizo-
pods the more easily does it stain with the aniline colours.
We may often see flagellated Monads taking up
filaments of Leptotrix several times as long as themselves
(Fig. 9), and finally enclosing them in their digestive
vacuoles. (Fig. 10.) It is sometimes possible to follow
1 Jenaische Zeitschrift, vol. xxiv., 1889, p. 109.
C 2
20
INFLAMMATION.
all the changes undergone by the bacteria within an
infusorium, as in the case of the diges-
tion by Stentor of the sulphobacterium
Thiocystis, observed by Le Dantec. 1
It is evident that the digestive func-
tion of the protoplasm of the Protozoa
must hinder the invasion of these
animals by the lower organisms, and
.it is only in certain special cases that
the latter can live as parasites within
the Rhizopoda and Infusoria. As I
have already mentioned an infective
disease of the former class of animals,
I will pass to the consideration of an
epidemic affecting the ciliated In-
fusoria.
In several species of Infusoria and
especially in Paramaecia, attention has
long been called to the presence in the
nucleus of a number of very fine rods,
which J. Muller, who first discovered
them, looked upon as spermatozoids.
These bodies were afterwards studied
by several observers, of whom I may
specially mention Balbiani and
Butschli, and were regarded by them
as parasitic bacteria. In reality they
are organisms quite distinct from
the Bacteriaceae, and belong to a
special group, consisting of several
species. Some of these develop in
the nucleus, replacing its contents,
1 " Recherches sur la digestion intracellulaire," Lille, 1891, p. 53.
Fig 9. A Monad in the
process of englobing a
filament of Leptotrix.
LECTURE II.
21
while another species attacks only the nucleolus.
(Fig. II.) The parasite, in its vegetative condition,
occurs in the form of elongated fusiform cells or rods
which multiply by transverse division or sometimes by
budding. Arrived at maturity, the parasites are trans-
formed into peculiarly shaped spores resembling in
their general appearance either bacilli or spirilla. 1
In spite of the abundance of these microbes in
organs so important as the nucleus and nucleolus, the
infected Infusoria remain capable of division, when
they do not die from exhaustion. In the process of
division a certain
number of para-
sites escape from
the nucleus into
the surround-
ing protoplasm,
whence they are
expelled just like
any other indiges-
tible body that has
Fig. io. Leptotrix enclosed in the digestive vacuole of a
been swallowed Monad.
by the infusorium. Hafkine has shown that, if placed
in exceptionally favourable conditions, the Paramaecium
may continue to divide and produce successive genera-
tions of infected infusoria ; at each division however,
the organism rids itself of a certain number of the
parasites, so that finally they may be all expelled and a
complete cure result.
Hafkine has never succeeded in producing infection
1 Vide the investigation of Hafkine, carried out at my instiga-
tion in my laboratory, and published in the Annales de FInstitut
Pasteur. Vol. iv., 1890, p. 148.
22
INFLAMMATION.
by introducing Paramaecia into capillary tubes contain-
ing spores of the parasite ; for the Infusorium, although
it swallowed a certain number of the spores, surrounded
them with a nutrient vacuole (Fig. 12, 13), and then
ejected them as it would any excrementitious matter.
In order that a spore should germinate, it must avoid
the digestive and expulsive
action of the protoplasm of
the Infusorium, and pene-
trate into the nucleus or
nucleolus, neither of which
has any digestive capacity.
We see that in this case,
as in the disease of the
Amoebae, the microbe, in order
to infect the Protozoon, has to
combat the power possessed
by its protoplasm of ejecting
or digesting the parasite.
The same holds good in all
f cases where we find the in-
Fig. n.-Paramscium with its nucle- Vader lodged in the digestive
olus filled with parasites.
contents of an infusorium.
The majority of the in-
fectious diseases of these Protozoa are doubtless
caused by the parasitism of suctorial Infusoria or
Acinetae, which I have already mentioned in the first
lecture. In spite of the delicacy of their cuticle,
these parasites offer complete resistance to the diges-
tive action of the protoplasm of their hosts, although
in many cases the latter (as e.g. Stylonychia) are
distinguished by their voracity and the ease with
which they digest their prey. As already mentioned,
o. mouth ; n. nucleus ; m. affected
nucleolus ; i>. c. contractile vesicle.
LECTURE II. 23
the young Acinetae fix themselves on to the surface
of other Infusoria, and penetrate into the endoplasm
of the latter by means of their active movements.
Once arrived in the central mass of protoplasm, the
parasites grow considerably, divide, and give rise to
a number (fifty or more) of young individuals, some
of which escape from the body of the infusorium to
attack another, after a cer-
tain period of freedom.
In order to survive in
the interior of the In-
fusoria, the Acinetae must
exercise some paralysing
influence on the digestive
function. It is probable
that these parasites secrete
Fig. 12. A Paramaecium which has taken
up some spores of the parasite.
a. b. c. d e. f. g. spores surrounded by a
vacuole ; . nucleus ; i'. c. contraccile
vesicle.
Fig. 13. A vacuole containing
spores (very highly magnified).
some toxic substance, since one often sees various Infu-
soria fall into a paralysed condition and die in con-
sequence of the attacks of free Acinetae. 1
By their growth within the Infusoria the Acinetae
give rise to degeneration of the nucleus, which breaks
up into small round granules. In many cases, however,
1 For an account of what is known about the Acinetae, see
Biitschli, " Protozoa," in Bronn's " Classen u. Ordnungen des
Thierreichs/ ; vol. iii., 1889, pp. 1823 and 1842.
34 INFLAMMATION.
these parasites do not kill their hosts, which may even
preserve their power of multiplication. Much more
dangerous for the unicellular organisms are the
infections produced by the fungi belonging to the group
of Chytridiaceae, which in most cases attack Infusoria
incapable of intracellular digestion, such as those which
obtain their nourishment simply by diffusion. They
may also attack Infusoria capable of intracellular diges-
tion, but in this case the infection occurs while the
Infusorium is in an inactive or encysted condition,
during which time digestion does not take place. The in-
tracellular Chytridium, after penetrating into the interior
of the Protozoon, becomes round and immobile, and
absorbs the substance of its host, which dies, while the
parasite gives rise to zoospores.
Fig. -n.Euglena viridis enclosing a Chytridium.
We will take as an example the Chytridium which so
often infects the Euglena viridis, and which was dis-
covered by Klebs. 1 Among these Flagellata, which
abound in stagnant water, we may find individuals
which are to all appearances in perfectly good health,
but enclose a round body, provided with a nucleus
and a very delicate cuticle. (Fig. 14.) This foreign
body grows gradually larger, and divides into a number
of small cells which become converted into conical
1 Untersuchungen aus d. botan. Institute in Tubingen, vol. i.,
1883. See also Hafkine, Annales des sciences naturelles : zoo-
logie, 1886, pp. 330, 336, &c.
LECTURE II.
z spores f
zoospores. (Fig. 15.) The zoospores bore their way
out of the Euglena and escape into the surrounding
water. In the course of this development, the affected
individual presents unmis-
takable signs of disease.
The green chromatophores
are rapidly absorbed, and
the Euglena becomes highly
anaemic. Its contents at
the same time undergo Flg<
pigmentary degeneration,
evidenced by the formation of scattered brown granules
which become gradually more and more numerous.
When the parasite has attained the zoospore stage, the
Euglena dies in consequence of the infection.
The encysted condition, during which the Euglena
viridis is protected by
a capsule, seems to
preserve it from the
attacks of the Chytri-
dium, since this is
only found within the
mobile Euglenae. On
the other hand, the
cysts of this infu-
sorium are often in-
Fig. i6.PaHdorina with one cell attacked by an vao - e d by \l\eP Olypha-
Olpidium. 7 t 1*1
gus Euglen<z> which
represents another genus of the Chytridiaceae.
The colony-forming Flagellata are equally subject to
infection by the Chytridiaceae. Pandorina morum (one
of the Volvocineae) is often attacked by an Olpidium.
The presence of the latter in the body of the flagellated
26
INFLAMMATION.
Monad causes a secretion of fluid, which collects to
form a vacuole. (Fig. 16.) The parasitic cell, which
is small and transparent, increases in size at the expense
of its host, and becomes filled with fatty granules
interspersed with transparent vacuoles. Soon after-
wards it sends out a conical process which pierces the
cell-wall of the Pandorina (Figs. 17, 18), and forms an
outlet by which the zoospores produced by the seg-
mentation of the contents of the parasite escape. At
Fig. 17. Another Pandorina with five infected cells.
other times the parasite, without giving rise to zoo-
spores, may secrete a thick external membrane, and be
transformed into a cyst.
As in the case of Euglena, the invaded cell undergoes
pigmentary degeneration, and always ends by dying
and disintegrating. The neighbouring cells of the
colony however remain quite unaffected. They pre-
serve complete mobility, show pulsation of their con-
tractile vesicles, and divide in a perfectly normal
manner. (Fig. 18.) The disease and death of one or
even of the greater number of the sixteen members of
LECTURE II. 27
the colony has absolutely no effect on the individual
cells which have escaped infection by the parasite.
This brief account of artificial lesions and of the in-
fectious diseases of the unicellular organisms, while
showing the inadequacy of our present knowledge on
the subject, at the same time enables us to appreciate
to some extent the general character of these phe-
nomena. With regard to those which are called forth
by the lesions, the most striking fact is the complete
power of regeneration possessed by these beings. As
Fig. 18. Part of an affected I'andorina; and the zoosporangium
of the parasite a.
we have seen, a detached segment can regain its normal
form in a very short time, some hours or even minutes
after the section.
After what has been said in the first chapter, we are
justified in assuming generally that the relations be-
tween the Protozoa and the micro-organisms which
infect them are to be regarded in the light of a struggle
between two living species. The parasites are often
nothing else than voracious organisms which, in conse-
quence of their minute size, do not attack their prey
directly, but make their way into the bodies of the
Protozoa which serve them for food. This carnivorous
28 INFLAMMATION.
nature of the parasites is seen not only in the Acinetae,
but also in the parasitic Flagellata which are allied to
the Vampirellae and other voracious organisms. In the
cases of infection however, the struggle assumes a more
complicated and indirect character. The parasite makes
its onslaught by secreting toxic or solvent substances,
and defends itself by paralysing the digestive and ex-
pulsive activity of its host ; while the latter exercises a
deleterious influence on the aggressor by digesting it
and turning it out of the body, and defends itself by the
secretions with which it surrounds itself.
Although these phenomena do not come under the
heading of the struggle for existence in the strictly
Darwinian sense (i.e. competition for the survival of
the fittest among individuals of the same species), yet
they are all more or less directly connected with the
struggle for survival that is always going on between
the representatives of the different orders of living
beings. In this struggle an important part devolves on
the power of intracellular digestion, which is so gene-
rally met with in the Rhizopoda and Infusoria and is
not entirely wanting even in the Protozoa which obtain
their food entirely by osmotic absorption.
LECTURE III.
Plasmodium of Myxomycetes Puncture by a glass tube Cauteri-
sation Chemical excitation Trophotropism Chemiotaxis
Habituation of the plasmodium Negative chemiotaxis
Repulsion of the plasmodium in the presence of bacteria
Digestion of bacteria by the plasmodium Sensibility of the
plasmodium Fixed plants having no true intracellular diges-
tion Necrosis and regeneration Waldenburg's experiments
Functions of the cell-wall De Bary's observations on
Peziza sclerotiorum Tumours of plants.
WE have now to consider the pathological phenomena
which occur in multicellular organisms ; and first of all
we meet with a group which is very important for
several reasons and especially interesting on account of
the simple organisation of its members. I refer to the
Myxomycetes a group presenting both animal and
vegetable properties and characterised by the fact that
it passes through a plasmodium stage. This plasmodium
is composed of the largest masses of protoplasm known
to occur in nature.
The plasmodium, as is well known, represents a colos-
sal amoeboid organism formed by the fusion of a large
number of zoospores of the Myxomycetes and enclosing
numerous nuclei embedded in a common protoplasm.
Branching in all directions, the plasmodium is able to
move about on the various objects (dead leaves, wood,
&c.) on which it lives ; it shows amoeboid movements
3O INFLAMMATION.
at the edges of its ectoplasm, while the inner layer of
its protoplasm (or endoplasm) is the seat of rapid
currents, recalling those of volcanic lava. The plas-
modium readily encloses any solid bodies within its
reach, and partially digests them by the help of a peptic
ferment and an acid which it secretes around its food. 1
The residue as well as the indigestible bodies are turned
out by the plasmodium, thus forming tracks which
mark the places where the protoplasmic processes have
been. At a certain time the plasmodium produces
Fig. 19. A part of the plasmodium enclosing a glass tube.
sporangia, which are usually in the form of minute
fruit, and enclose a number of spores provided with a
tough envelope.
On account of its great extent, which may amount to
as much as a foot or more, the plasmodium offers many
advantages for the study of protoplasm in general and
of pathological phenomena in particular.
In order to observe the effect of a lesion upon a plas-
modium, we may introduce a solid foreign body, such
1 The peptic ferment was discovered by Krukenberg. Unter-
suchungen aus dem physiol. Instit. d. Univ. Heidelberg, vol. ii.,
1878, p. 273. Concerning the acid of the plasmodia, see Annales
de flmtitut Pasteur, 1889, p. 25.
LECTURE III.
as a minute glass tube, into the protoplasm of a Phy-
sarum. The puncture by the tube tears a part of the
plasmodium, which diffuses into the surrounding fluid.
But the chief mass of protoplasm is in no way affected
by the tube, which it moreover englobes after a short time
just as if it were a particle of food. (Fig. 19.) After
retaining the tube for a longer or shorter time, the
plasmodium ejects it like any other substance that it is
unable to utilise as nourishment.
We may irritate the
plasmodium in another
way. If we take a
specimen (such as the
yellow plasmodium of
Physarum) on an object
glass and touch its
central part with a
minute glass rod pre-
viously heated in a
flame, we shall produce
thermal excitation in-
stead of a mechanical lesion. Immediately after being
touched, the central part of the plasmodium dies and
may be clearly distinguished from the living peripheral
portions, which remain motionless as if nothing had
occurred, and are unaffected by the necrosed portion. A
few hours later however the plasmodium wakes from its
passive condition and creeps away from the dead part.
Chemical irritants operate in a still more powerful
way. If we apply a minute fragment of nitrate of
silver to the edge of a plasmodium of Physarum lying
on a piece of glass, and wash the injured spot directly
afterwards with a one per cent, solution of sodium
Fig 20. Plasmodium cauterised by nitrate of
silver.
32 INFLAMMATION.
chloride, (in order to precipitate any nitrate which may
have become dissolved,) we shall see that the edge
touched by the nitrate of silver dies and becomes
detached from the rest of the plasmodium. (Fig. 20.)
The latter reacts immediately by a rapid change in the
direction of its movements. Before the operation, the pro-
toplasmic currents were tending towards the edge where
the nitrate was applied (this spot having been chosen
for the experiment on this account) ; directly afterwards
Fig. 21. The same plasmodium 50 minutes after the stage
represented in Fig. to.
they were turned towards the sides of the plasmodium
and soon assumed a direction completely opposed to
the original one. (Fig 21.) At the end of an hour
from the beginning of the experiment, the plasmodium
had moved some distance from its first position, leaving
the dead residue behind.
In both of these experiments the plasmodium for-
sakes the portions injured by thermal or chemical
agencies. The much more rapid removal of the plas-
modium in the second case may be attributed to the
more powerful influence of the nitrate of silver.
LECTURE III. 33
We thus see that irritating agents excite in the plas-
modium either a course of events similar to those which
accompany the taking up of any solid nutriment, or else
a more or less marked repulsion. In attempting to
produce a reaction which should correspond to the con-
dition of inflammation in the higher animals, we have
brought about the phenomena of attraction or repulsion
which occur so frequently in the lives of plasmodia and
the inferior animals generally.
As early as 1884, Stahl ' discovered that a decoction
of dead leaves (on which so many of the Myxomycetes
feed) has the property of attracting the plasmodia,
whereas solutions of salt, sugar, and numerous other
materials, act in a contrary manner, repelling the
plasmodia to a greater or less distance. In connecting
these phenomena with those of nutrition, Stahl gave
them the name of positive trophotropism when it was a
question of attraction, and of negative trophotropism in
cases of repulsion. Pfeffer, 2 having found that the female
organs of certain cryptogams (ferns, mosses, and
selaginellae) attract the spermatozoids with a different
object from that of nutrition, grouped all these various
forms of sensibility to chemical agents under the general
heading of chemiotaxis (positive or negative), a term
which was very soon generally adopted.
Since, as we have just shown, these phenomena of
sensibility play an undoubted part in pathological pro-
cesses, it is desirable to consider them in somewhat
greater detail. Chemiotactic phenomena are met with
not only in the Myxomycetes and the spermatozoids of
1 Botanische Zeitung, 1884, Nos. 10-12.
2 Untersuchungen aus d. botan. Institute in Tubingen, vol. i. p.
363-
D
34 INFLAMMATION.
the above-mentioned cryptogams, but also in Bacteria,
Flagellata, Volvocineae, 1 and in the zoospores of fungi,
such as the Saprolegniaceae. 2 Hence it is evident that
we have here to do with a phenomenon of general
import.
It cannot be denied that by means of positive chemio-
taxis the organisms are directed in their search after
nutrient substances, and are enabled to approach the
bodies with which they have to establish relations, as
in the case of the spermatozoids attracted by an ovum.
By means of negative chemiotaxis, on the other hand,
they can escape injurious influences. This rule,
although true generally, cannot be applied to every
particular case. Thus Pfeffer 3 has seen spirilla and
animalcules (' bodons ') dart into too highly concentrated
solutions of sugar and glycerine, to which attracting sub-
stances had been added, where they inevitably perished.
The analogy between these phenomena and the sen-
sations of man and the higher animals is obvious. One
among many proofs of this analogy consists in the fact
that the chemiotaxis of the lower animals is subject to
the same law of Weber which has been established for
the sense perceptions of man. In order that a bacterium
(B. termo} or the spermatozoids of ferns (the organisms
on which Pfeffer has made his remarkable observations)
may be affected by a change in their surrounding media,
it is essential that this change should attain a certain
degree. Thus the Bacterium termo placed in a solution
of peptone of definite strength, will not move towards a
1 Untersuchungen aus d. botan. Inst. in Tubingen, vol. ii. 1888,
p 582.
2 Botani 'sche Zeittmg, 1890, Nos. 7-11.
3 Unters. a. d. botan. Inst. in Tubingen, vol. ii. p. 627.
LECTURE III. 35
more concentrated solution of peptone, until this is five
times as strong as the first solution. Having ascertained
these proportions, Pfeffer formulated for the chemiotaxis
of these unicellular bodies the same law as for the sense
perceptions of man, viz. that when the excitation is
increased in geometrical proportion, the sensation is in-
creased in arithmetical proportion, or, in other words, that
the reaction varies as the logarithm of the excitation.
Now although the two orders of phenomena conform
to the same law, there is quantitatively a great difference
between them. Man can appreciate a difference of
weight equalling one-third, of temperature amounting to
one-thirtieth, of light equivalent to one-hundredth, where-
as the spermatozoids of ferns are not affected by and do
not react to an alteration in the chemical composition,
until the quantity of the substance which is acting on
them is augmented twenty-nine times. The Bacterium
termo is indifferent to an increase of concentration until
this amounts to four times the original concentration. 1
In order to test the chemiotactic sensibility of the
plasmodium, I placed several specimens of the plasmo-
dium of Didymium farinaceum in O'l, O'OI, 0*05, 0*005,
and 0*0005 per cent, solutions of hydrochlorate of
quinine. The last two solutions did not prevent the
plasmodium from advancing or even from inserting
several processes into them ; whereas the first three
solutions exercised a pronounced negative chemiotactic
effect. (Plate II., figs. 3-6.) The plasmodium therefore
can appreciate the difference between 0^05 and 0x305
per cent, of hydrochlorate of quinine.
The plasmodium, in common with other of the lower
organisms, has the important power of growing gradually
1 Pfeffer, loc. tit., vol. ii. p. 637.
D 2
36 INFLAMMATION.
accustomed to solutions which in the beginning it
avoided. Stahl was the first to notice that the plasmo-
dium of Fuligo, which is at first repelled by a solution
of sea-salt in the proportion of 2 per cent, or less,
after having been deprived for some time of water
finally adapts itself to the changed conditions and dips
its processes into the salt-water. Here we have an
instance of negative chemiotaxis which, as the result of
imperceptible alterations in the protoplasm, is converted
into positive chemiotaxis.
As this fact is of great importance from a general
point of view, I was desirous of watching the process
myself. For this purpose, I placed a plasmodium of
Physarum extended on a glass slip into a vessel con-
taining a solution of 0*5 per cent, sodium chloride.
The plasmodium immediately showed negative chemio-
taxis and moved away from the surface of the liquid.
It was then changed to another vessel containing a 0*25
per cent, solution of the same salt. The plasmodium
was at first repelled, but after the lapse of a few hours
it drew near the liquid into which it then immersed the
end of its processes. With the view of noting how far
this power of adaptation extended, I replaced the
plasmodium in the first vessel with the 0*5 per cent,
salt solution. It again receded from the fluid ; but at
the end of about twelve hours it approached the surface
of the water, without however touching it.
By means of negative chemiotaxis, therefore, the
plasmodium is enabled to avoid injurious influences ;
as we have seen, it recedes from bodies which burn it,
such as the nitrate of silver, and even from necrosed
portions of its own organism, as in the experiment of
the application of a heated object. It is probable that
LECTURE III. 37
the same property may serve to protect the plasmodium
against the attack of other organisms, especially of
pathogenic microbes.
Stahl has observed that plasmodia are never attacked
by parasites. This fact he seeks to explain by referring
it to the facility with which the plasmodia move about
from place to place, as well as to their power of
expelling foreign substances a property which is con-
nected with the intracellular digestion of solid bodies.
Although no direct observations on the expulsion of
parasitic organisms by the plasmodium have so far been
made, yet it is extremely probable that this occurs,
especially as Pfeffer 1 has watched the plasmodia of
Chondrioderma ejecting living Pandorinae and Diatoms.
Moreover the direct observations on the expulsion of
parasitic spores by the Paramaecia tend to support
Stahl's deduction on this subject.
The following experiment was made with the object
of ascertaining the true significance of the movements
of the plasmodium. I spread out a plasmodium of
Physarum on a slide, and placed it midway between
two small glass vessels, one of which was filled with a
stagnant infusion of dried leaves, full of bacteria,
infusoria and other of the lower organisms, the other
with the same infusion after filtration through several
layers of filter-paper. The two ends of the plasmodium
were connected with the liquid in the two vessels by
strips of blotting-paper. The plasmodium soon began
to approach the filtered liquid, moving along the strip
which was soaked in it. Another experiment made in
1 " Ueber Aufnahme und Ausgabe ungeloster Korper." Ab-
handlungen d. math. phys. Klasse der k. sacks Gesellsch. d. Wissen-
schaften^ vol. xvi. 1890, p. 161.
38 INFLAMMATION.
the same way, with a few slight modifications, was
attended by exactly the same result, showing that the
plasmodium preferred the liquid which was free from
microbes. In order to ascertain how far this preference
extended, I repeated the experiment, only exchanging
the filtered fluid for a fresh infusion of dead leaves in
cold water, which was consequently colourless. This
time the plasmodium advanced towards the stagnant
infusion, in spite of the microbes which it contained.
The repulsion of the plasmodium in the presence of
the lower organisms is evidently merely relative ; this
is in harmony with the fact that the Myxomycetes in
their amoeboid condition have the power of englobing
microbes. Saville Kent has observed amoeboid
zoospores of Physarum lussilaginis which were filled
with bacteria. Later on Lister 1 made some highly
interesting researches on the inclusion of bacteria by
the zoospores of different Myxomycetes. The bacteria,
seized on by the pseudopodia, are dragged into the
interior of the amoeboid plasma and lodged in the
nutritive vacuoles. Here they become less and less
distinct, and appear to be almost completely dissolved.
A zoospore of Chondrioderma difforme will completely
digest a couple of large bacilli in about an hour and a
half.
The powers of digestion and expulsion possessed by
the plasmodium, combined with its property of negative
chemiotaxis, are of real service to it in reacting against
injurious excitants.
The plasmodia, in common with a considerable num-
ber of other lowly organisms, are endowed with several
forms of sensibility besides that known as chemiotaxis.
1 Journ. of the Linncean Soc., Botany,vo\. xxv. 1890, p. 435.
LECTURE III. 39
They avoid sunlight, and are powerfully attracted to the
damper places, thus evincing a kind of hydrotropism.
Moreover, this positive hydrotropism is converted into
negative hydrotropism just before the period of fructifica-
tion, when the plasmodium seeks a dryer spot (Stahl).
The plasmodia are also endowed with tactile sensi-
bility.
Fixed multicellular plants, which have no powers of
locomotion or of digesting and excreting foreign bodies,
react to the various injurious factors with which they
may come in contact in a different manner from the
plasmodia, which can either avoid dangerous objects
or can eliminate them by digestive or excretory pro-
cesses.
A thorn introduced into the plasmodium is dealt with
like any other foreign body that becomes enclosed in the
amoeboid mass. As it cannot be digested, it is simply
ejected altogether. If introduced into the tissue of a
plant, the same thorn would produce a lesion of the cells,
and the injured cells would inevitably perish. The
damage is soon repaired with the help of the neighbour-
ing cells, which rapidly multiply and form either a mass
of cork or a true scar composed of several tissues. 1 In
both cases there is active cellular neo-formation at the
injured spot. Waldenburg, 2 who has studied these
phenomena from the point of view of their analogies
with inflammation in the higher animals, says : " Plants
may therefore suffer from inflammation, if by this
term we imply only the lesions produced by the irri-
tant, as well as the tumour induced by these lesions,
1 Frank, " Die Krankheiten der Pflanzen." Breslau, 1880, vol. i.
p. 95, etc.
2 VirchoiiJs Archiv, vol. xxvi. 1863, pp. 145 and 322, Tab. v.
4O INFLAMMATION.
and leave out of account the blood-vessels and nerves "
(p. 344). Inflammation would thus be only an irrita-
tion of the tissues (tumefaction, growth) plus a vascular
congestion.
The phenomena of repair in plants have on more
than one occasion been quoted in support of the
attraction theory of inflammation, and especially in
favour of Virchow's theory of a nutritive and formative
hyperactivity of the inflamed tissues. But unfortu-
nately no account has been taken of the conditions
intermediate between plants and the higher animals,
and consequently the most characteristic phenomena of
true inflammation have been lost sight of altogether.
The new cells produced at the seat of injury in
plants frequently secrete thicker and tougher cell-
walls than usual. The cuticle in fact represents the
true protective organ of the plant. The instance
quoted in the preceding lecture of a Chytridium which
only attacks the Euglena in its mobile and never in its
encysted condition, may be taken as a confirmation of
this statement. The walls of the vegetable cells are
too resistant for many of the microbes, and especially
for those which cannot penetrate actively into the
tell-contents. This is probably the reason why
bacterial infection so rarely takes place in plants. On
the other hand, vegetable organisms are very liable to
be attacked by fungi, which grow rapidly and in many
cases secrete a diastase which dissolves the cellulose
cuticle of the plant. The parasitic fungus, once within
the cell, absorbs the contents without further hindrance.
The cells that are thus invaded die, while the others
that remain alive, undergo rapid proliferation which
results in the overgrowth of parts, amounting in some
LECTURE III. 41
cases to a hypertrophy of the entire organism affected,
as in the case of Euphorbia Cyparissias when infested
by the aecidium of Uromyces Pisi. The presence of a
parasite, whether it be a fungus or a member of the
animal kingdom, often excites the formation of special
tumours or galls.
As in the healing of wounds, infections l in plants are
accompanied with regenerative phenomena, dependent
on the rapid proliferation of the cells that are not directly
affected. We do not however find processes analo-
gous to the essential phenomena of inflammation in the
vegetable world. For these we must pass to the con-
sideration of the representatives of the animal kingdom.
1 The most carefully recorded instance is undoubtedly that of
the Peziza sclerotiorum on which De Bary {Botanische Zeitung,
1886) made his classical observations. This fungus germinates on
the surface of the plant and sends out filaments with which it
subsequently penetrates the tissues. (The Peziza sclerotiorum
invades a large number of plants.) In order to effect an entry,
the parasitic filaments secrete oxalic acid and a ferment which
dissolves the cellulose. The parasite feeds on the juice of the cells
which have perished under the influence of its secretions, and its
mycelium grows into the interstices between the cells, rarely
penetrating the latter. De Bary has observed that the Peziza
easily finds its way into young plants, but is unable to force an
entrance into older plants of the same species. This immunity is
probably owing to the fact that the parasite cannot dissolve the
cellulose of the old cells. The control experiments showed in fact
that the extracted juice of the fungus was able readily to digest the
walls of the young cells and was quite without effect on those of
the older individuals of the same plant.
It is evident that the power of resistance possessed by the
vegetable cell is specially dependent on the toughness of its
cuticle. The parasite in order to infect the plant must first
perforate or dissolve this membrane.
LECTURE IV.
Transition from the unicellular organisms to the Metazoa Sketch
of the phagocytella theory Protospongia Sponges : their
organisation their three layers their nutrition Intracellular
digestion Ablation of parts of the Sponge Artificial division
Introduction of pointed bodies Utilisation of foreign bodies
to assist in forming the skeleton Fate of the organisms which
have penetrated into the interior of the Sponges Protective
function of the ectoderm Comparison with the Myxomycetes
Comparison with the inflammation of vertebrata.
IN passing to the animal kingdom, we have to confess
that we are at present ignorant of the way in which the
multicellular animals or Metazoa are derived from the
Protozoa. The gap between the most highly developed
members of the latter class and the lower kinds of
Metazoa is too wide and can only be bridged over by
hypotheses based on the embryological study of different
animals. Putting aside several groups of parasites
which have undoubtedly lost much of their primitive
character (Dycyemides, Orthonectides) , we find that even
the simplest forms of the Metazoa, such as the Sponges,
are composed of a multiplicity of organs, arranged in
three well-known layers : the ectoderm, the mesoderm
and the endoderm. In order to obtain a knowledge of a
more simple condition of animal life, we must turn to the
embryos of Sponges and of other inferior organisms, such
as the Medusae and their allies. Here we may readily
LECTURE IV. 43
meet with stages in which the animals are composed of
two layers, one of which constitutes an enveloping
membrane for the larva, the other is formed by the inner
cells, grouped in a different manner. These latter cells are
sometimes collected into a solid mass, a kind of paren-
chyma, composed of amo3boid elements ; or they may
be arranged evenly to form an epithelial layer sur-
rounding a digestive cavity. The question as to which
of these two forms may be considered the more primi-
tive has been much discussed. I think that the
absence of a digestive cavity, the irregular shape of the
cells, together with various other facts afforded by the
comparison of the embryogeny of many lower animals
(which do not come within the range of this pathological
survey), lead to the conclusion that the parenchymatous
stage is the more primitive of the two. I have called
this stage by the name of Phagocytella^ on account of
the power which the cells of the inner layer possess of
englobing various solid bodies, and more especially for
the reason that the digestive cells of the whole organism
are derived from this layer. The latter in the first
place produces the endoderm which lines the intestinal
canal and its appendages, and secondly the whole or a
part of the mesoderm which also includes a large
number of digestive cells or phagocytes.
The phagocytella stage may readily pass into the
Gastrula stage, 2 which has two epithelial layers, one
of which represents the wall of the primitive intes-
tine that opens by a primordial orifice or blastopore.
1 Cf. my account of the Phagocytella theory in my " Embryolo-
gische Studien an Medusen," Wien, 1886.
8 For an account of the Gastraea theory, see Hasckel " Gastraea-
Theorie." Jena, 1874.
44
INFLAMMATION.
This gastrula is so to speak the starting-point for all
the Metazoa.
The structure of the Metazoa when reduced to its
simplest condition, that of the phagocytella, is some-
what analogous to that of certain colony-forming
Protozoa. In these cases the colonies are made up of
two kinds of individuals : the flagellated individuals
Fig. 22. Protospongia Heeckelii (after Saville Kent).
forming a sort of outer layer (Fig. 22), and the
amoeboid individuals, situated in the inner mass of the
colony. The former would thus correspond to the
elements of the ectoderm, which so frequently consists
of flagellated cells, and the latter would constitute a
kind of inner parenchyma, composed of amoeboid cells,
which are at the same time phagocytic in their nature.
In these colonies of infusoria, termed Protospongia by
LECTURE IV. 45
their discoverer Saville Kent, 1 the two layers are not
yet clearly defined, since the individuals of which they
are made up pass readily from the one form into the
other.
By means of these colonies on the one hand and of
the organisms resembling the phagocytella on the
other, we are almost in a position to bridge over the
gap between the Protozoa and the Metazoa.
I should not have dwelt upon these hypotheses in
these lectures on the comparative pathology of inflam-
mation, did not their consideration afford an opportunity
of studying the general significance of the presence of
these amoeboid cells which are able to englobe solid
bodies. We have met with them in the various classes
of Protozoa, and we find them again even in the most
primitive forms of Metazoa. Now the co-operation of
amoeboid cells in inflammation as it occurs in the
vertebrates is a fact of the greatest importance, which
has received almost universal acceptance.
As low down in the animal scale as the most inferior
Metazoa we have to do with these cells. The Sponges
are of such undifferentiated organisation that they were
long considered to be colonies of Protozoa, consisting,
like the Protospongia, of separate flagellated and
amoeboid individuals. Later on it was however ascer-
tained that they bore a certain relationship to the
Polyps and their allies (Ccelenterata). It was then
found that they are formed of three characteristic layers.
The outer layer or ectoderm covers the whole structure
with flat epithelial cells, the contours of which are very
clearly defined after the application of a solution of
nitrate of silver. The cells themselves are evidently
1 " The Manual of Infusoria," 1880-1882.
46 INFLAMMATION.
contractile a property which is more readily observed
at the free edges of the young cells, where amoeboid
prolongations belonging to the ectodermic elements are
seen. The contractility of these cells is certainly con-
cerned in the remarkable phenomenon of the opening
of the numerous pores which are scattered over the
whole surface of the Sponge and are bordered by two or
more flat cells. These pores open to permit of the
passage of a stream of water with the minute particles
which it holds in suspension. The liquid first enters
a system of efferent canals which are also lined by a
pavement epithelium, the origin of which is not yet
known. It then passes into canals or into round cavities
(the ' ciliated chambers ') which are covered with a cylin-
drical epithelial layer, the cells of which possess a single
large flagellum. These cells, which present a striking
analogy to many flagellated infusoria, form part of the
endoderm and represent typical phagocytes, since they
attract and englobe a large number of fine granules
which are carried along by the stream.
Besides these flagellated phagocytes of endodermic
origin, the Sponges contain a large number of mobile
cells, which are typical amoebae and form part of the
mesoderm, being situated between the ectoderm and
the cylindrical epithelium. Although it has not so far
been definitely ascertained how the foreign particles
penetrate the mesoderm after they have reached the
interior of the Sponge, yet it has been clearly shown
that they are largely absorbed by the mesodermic cells
themselves. If a coloured substance, such as carmine
indigo or sepia, be added to the water in which the
Sponges are immersed, it will be found soon afterwards
that many of the grains of colouring matter have been
LECTURE IV. 47
enclosed by the endodermic cells, as well as by the
amoeboid phagocytes of the mesoderm.
In certain Sponges (as for instance in several cal-
careous Sponges,) there are very few mesodermic cells,
which consequently take but a small part in englobing
foreign bodies ; in others again, especially in the
silicious kinds, the mesoderm is much better developed,
and its more numerous cells can therefore take in a
proportionately larger number of these minute particles.
There are a few species, such as the Siphonochalina
coriacea, whose mesodermic cells alone enclose all
foreign bodies, so that the cylindrical cells of the
endoderm merely serve to keep up the continuous
passage of the fluid through the Sponge. The phago-
cytes of both layers have the power of rejecting in-
soluble matters, which collect in the larger efferent
canals and are expelled from the organism through
large apertures of crater-like shape, the walls of which,
according to some authors, are furnished with muscular
fibres.
We are however chiefly concerned here with the fact
that the mesodermic phagocytes are able to digest the
substances as well as to englobe them, and to reject the
insoluble residue. Many years ago Lieberkiihn * ob-
served the digestion of Infusoria which had found their
way into the mass of amoeboid cells of soft-water
Sponges, and pointed out the analogy of this pheno-
menon with the digestion of Infusoria by the Rhizopoda
or other Protozoa. This has been confirmed by
other investigators. Thus I have seen 2 Oxytricha,
Glaucomae, and Actinophrys dissolved in the midst of
1 Mailer's Archiv fur Anat. und Fhys., 1857, p. 385.
2 Zeitschr. f. wissench. Zoologie, vol. xxxii. 1879, P- 37 !
4 8
INFLAMMATION.
a mass of the mesodermic phagocytes of a young
Spongilla, and afterwards the foreign bodies which had
been swallowed by these Protozoa englobed by the
same phagocytes. The Euglenae, when sucked into the
SpongillaB by the current, become likewise surrounded
by the mesodermic phagocytes ; but whereas their
protoplasm is then digested, the grains of chlorophyll
and paramylum remain intact for an indefinite time.
Viauari.
Fig. 23. A mesodermic phagocyte of a young Spongilla surrounded by several
ectodermic cells.
The mesodermic cells of young Spongillae, immedi-
ately after their escape from the gemmules, can enclose
foreign bodies even before the endoderm is developed.
The young Sponge at this stage consists only of a layer
of flat ectodermic cells and an irregular mass of mesoder-
mic cells, a certain number of which soon begin to
secrete spicules. If carmine grains be placed in the
LECTURE IV. 49
water inhabited by the Spongillae, they find their way in
without apparently injuring the wall, and are at once
englobed by the amoeboid phagocytes of the mesoderm.
(Fig- 23.)
The above facts are so invariable in their occurrence
and may be so readily observed, that it is very extra-
ordinary that de Lendenfeld, 1 in his monograph on the
physiology of Sponges, should seek to throw doubt on
the matter. This author asserts that carmine, when
added to the water which filters through the Sponges, is
very rarely deposited in the amoeboid cells and then
only where there is some lesion of the outer layer of
cells, and that in a normal Sponge it is only the
cylindrical cells of the endoderm which will seize on the
carmine. De Lendenfeld lays stress on these con-
clusions, in spite of the fact, which he acknowledges,
that the fat-globules of milk are readily absorbed by
the mesodermic phagocytes. This is a sufficient proof
of the part taken by these cells in the intracellular
digestion of Sponges. Moreover his memoir con-
tains direct indications of the presence of carmine
grains in the amoeboid cells of the mesoderm. Thus
this author describes that he has seen numerous carmine
grains in the phagocytes of Chondrosia reniformis, the
species on which he has bestowed the most study.
Further, he has even found them in these cells two
hours and a half after the introduction of the carmine
into the water, at a period when the cylindrical elements
of the chambers no longer contained any granules.
Since these facts remove all doubts that the amce-
1 " Experimentelle Untersuchungen iiber die Physiologic der
Spongien," Zeitschr. /. wissensch. Zoologie, vol. xlviii. 1889, p.
406.
E
5O INFLAMMATION.
boid cells of the mesoderm of the Spongillae have the
power of taking in and digesting foreign particles, I
have endeavoured to ascertain the conditions which
might influence this function. Following up the know-
ledge we have acquired concerning the typical in-
tracellular digestion in the Protozoa and the Myxomy-
cetes, which secrete around the object they have en-
globed an amount of acid sufficient to convert the
colour of litmus from blue to red, I placed a few grains
of blue litmus into the water containing some young
Fig- 24. A glass tube surrounded by the mesodermic phagocytes of the Spongilla.
Spongillae which had developed from gemmules. Like
most minute particles suspended in the water, these
grains were soon englobed by the Sponges and were
found to have been chiefly taken up by the mesodermic
phagocytes. The litmus however did not change
colour, even after a prolonged stay in the cells ; hence
it is evident that the digestion of the Spongillae
cannot proceed in an acid medium. This fact is in
complete harmony with Krukenberg's discovery l of a
tryptic ferment in the glycerine extract of several
varieties of Sponges.
1 " Grundziige einer vergleichenden Physiologic der Verdauung."
Heidelberg, 1882, p. 52.
LECTURE IV. 51
If now we introduce a sharp substance, such as a
minute glass tube or a spicule of asbestos, into a Sponge,
the greater part of it will be situated in the mesoderm,
where it will be in immediate proximity to the amoe-
boid cells. The latter surround the foreign body either
partially or completely, i.e. they react as if the sub-
stance were merely a nutritious body of larger size than
Fig. 25. Vegetable filament surrounded by the phagocytes of a Spongilla.
usual (Fig. 24). Sometimes the cells accumulate very
sparsely or not at all round the article introduced,
showing that too weak a reaction has been excited.
On other occasions inert bodies, such as vegetable
filaments, will attract a considerable number of phago-
cytes which surround them and become partially fused
into small plasmodia (Fig. 25).
In some members of this group the grains of sand
and of other hard materials, which they happen to have
E 2
52 INFLAMMATION.
taken up, become surrounded with a mass of spongine,
secreted by the mesodermic cells. In such cases the
foreign particles are utilised by the Sponge to increase
the solidity of its skeleton.
As I have remarked above, the mesodermic cells can
also enclose living organisms which have penetrated
into the interior of the Sponge, where they subse-
quently undergo digestion by the phagocytes. Organ-
rig. 26. Leptotrix surrounded by the phagocytes of the Spongilla.
isms of greater resisting powers may escape this fate
and may remain for a shorter or longer time within the
body cavity of the Sponge without undergoing any
alteration whatever. Thus I have seen filaments of
Leptotrix living uninjured inside the mesoderm of
young Spongillae, and surrounded by a plasmodium
formed by the fusion of a number of mesodermic cells
(Fig. 26). In certain Sponges (Hircinia echinata and
Ceraochalina gibbosd) Keller has found the eggs of
Annelida and Crustacea developing undisturbed in the
LECTURE IV.
53
mesoderm, and surrounded by masses of amoeboid
cells which formed a regular follicle round them (Fig.
27).
These facts show that any foreign body, which has
by some means or other reached the parenchyma of
the Sponges, excites the mesodermic phagocytes, which
either englobe them, or collect in a mass or even fuse
together in large numbers round them. If the foreign
bodies are easy to digest, they are soon dissolved ; if
however they prove resistant, they remain in the in-
Fig. 27. Crustacean ova surrounded by the phagocytes of the Ceraospontia,
(After Keller.)
terior ot the Sponge, surrounded by the cells. This
phenomenon, which frequently occurs among the
Spongiaria, may be regarded as an instance of a kind
of commensalism. These soft cellular organisms,
being readily penetrated, are very suitable as an abode
for many aquatic animals on account of the continual
stream of water which passes through them, bringing
nourishment to their guests. Hence there are a large
variety of the latter, from the Algae (Zoochlorella and
Zooxantellcs] which inhabit the interior of the meso-
dermic cells, to the Polypi (Stephanoscyphus], Annelida
and Crustacea, which take refuge in the canals and
54 INFLAMMATION.
parenchyma of the Sponges. So far, real parasites,
with the infectious diseases they bring in their train,
have not been discovered. It may be that the phago-
cytes are so effective as to utterly destroy the microbes
which enter the Sponges, or it may be that our know-
ledge is incomplete on the subject.
If now we consider the analogy existing between the
Sponges on the one hand and the Protozoa and Myxo-
mycetes on the other, we find that the digestive and
excretory functions take a prominent part in the re-
action against the foreign bodies which might injure
these organisms. In the case of a sponge or of a plas-
modium, this reaction consists merely in the enclosure
of the particles, and their subsequent digestion, if they
are digestible, or in their expulsion from the body, if
this is not the case. In the Myxomycetes this function
is performed by the whole of the protoplasmic contents,
whereas in the Sponges it is confined chiefly to the
mesoderm, though the endoderm also takes some part.
The outer covering or the ectoderm generally is however
not unconcerned in the work of protecting the organism
against all manner of injurious agents. The flat cells
of which the ectodermic layer consists, are contractile
and sensitive ; by their contraction the pores are opened
and allow the surrounding water to pass into the interior
of the sponge, so long as this does not carry any
harmful products along with it. It has long been
known that in order to follow the mode by which the
coloured granules penetrate into the cells to the best
advantage, experiments should be made with starving
specimens. As soon as the sponge is sufficiently filled
with the minute particles which have been brought by
the water, the pores refuse to allow the passage of any
more by remaining closed.
LECTURE IV. 55
According to de Lendenfeld, the Sponges do not
open their pores to harmful substances, whether these
are in solution or consist of granules suspended in the
water. Of all the bodies which he made use of, such as
carmine, starch, milk, the last was the only one that at
once found its way inside the sponge ; in the case of
carmine, the pores at first remained shut, although they
opened after a short while. The solutions of different
toxic substances, as morphine, veratrine, or strychnine,
caused contraction of the pores, which did not relax for
some little time.
There is an analogy as well as a difference between
the mode of action of the ectodermic cells, which are
contractile although not phagocytic, and that of the
plasmodia of the Myxomycetes. The analogy consists
in a sensibility towards the chemical composition of the
surrounding medium ; the difference is shown in the
manner of reaction. The mobile cellular colony the
plasmodium is repelled from the object which calls
forth the exhibition of its sensibility (negative chemio-
taxis, thermotaxis or otherwise) ; the motionless organ-
ism the Sponge avoids the same object by refusing
entrance to it.
In spite of the limitations of our knowledge, we are
in a position to affirm that in their struggle against
various injurious bodies, the Sponges make use of all
their cellular properties, more especially of the sensi-
bility and contractility possessed by the ectodermic
elements, and the power of englobing and digesting
common to mesodermic and endodermic cells. These
results may serve as a starting-point for the inquiry
into the more complex phenomena of reaction in other
animals.
LECTURE V.
Ccelenterata, Echinodermata and Vermes Traumatism and re-
generation in Hydra Accumulation of phagocytes in Acalepha
(Scyphomedusae) Phagocytes of star-fishes Inflammation in
Bipinnaria Reactive changes in the perivisceral cells of the
Annelida Phagocytic reaction in the diseases of Nais and
Lumbricus Struggle between the phagocytes of Lumbricus
and Rhabditis Microbic infections of Worms.
ALTHOUGH the Ccelenterata are distinguished from the
Sponges by their higher organisation, yet there are a
number of members of this group which consist of
only two layers of cells, the mesoderm being completely
absent. Since it is the mesoderm which, as we have
seen in the Sponges, plays the principal part in patho-
logical processes, it would be interesting to know how
those processes are carried out in such animals as Hydra
and its congeners, which have only two layers of cells.
As far back as the last century the phenomena
evoked in the fresh-water Polyps by every kind of
injury were often the subject of observation. Trembley
first pointed out the astonishing power of regeneration
possessed by this organism. A Hydra may be cut up
into several pieces, pierced with pointed bodies, and in
general maltreated to an extraordinary extent without
preventing a speedy and complete reintegration. In
one experiment of Ischikawa ' the front part of a
1 Zeitschr. f. wissensch. Zoologie, vol. xlix, 1889, p. 433.
LECTURE V.
57
Hydra had completely recovered within twenty minutes
after the infliction of an injury. Hydrae, cut in two
longitudinally, and stretched out on a cork, are able to
grow again into complete animals in little more than
twenty-four hours.
In another experiment Ischikawa cut off the head
and tentacles of a Hydra, and made a longitudinal slit
down the trunk, which he fixed on a piece of cork in
such a manner that the endoderm was directed out-
wards. In order to injure this layer the hydra was
Fig. 28. Regeneration of a Hydra (after Ischikawa).
taken out of the water in this position, and exposed to
the air for five minutes. On then detaching it from the
cork and putting it into the water, the Hydra first rolled
itself up into a cylinder, the outer surface of which was
formed by the endoderm (Fig. 28, i) ; soon, however, it
rolled itself in the opposite direction, so that the two
layers occupied their normal relative positions. During
this inversion, however, the experimenter had inserted
a filament of an alga between the edges of the cut, so that
they could not fuse together (Fig. 28, 2) . The Hydra then
changed its position, and finally grew into a closed sac
(Fig. 28, 3) which acquired a mouth and tentacles and
58 INFLAMMATION.
formed a perfect Hydra (Fig. 28, 4, 5) within six days
after the commencement of the experiment.
Punctures and other artificial lesions heal with extra-
ordinary rapidity, without any accumulation of phago-
cytes at the injured spot. But although no accumula-
tion takes place owing to the absence of a mesoderm,
yet we must not imagine that the phagocytic function
is completely wanting in these animals. The whole of
the endoderm in the Hydra consists of stationary phago-
cytes in the form of epithelial cells, which are capable
of putting out amosboid processes from their free sur-
face, and ingesting various foreign bodies.
In the marine colony-forming Hydromedusae, not
only the endoderm, but occasionally the ectoderm as
well, consists of phagocytes which are of great import-
ance on account of their prophylactic action. 1 These
creatures like the Hydra have the power of regeneration.
If the head of one of these Hydrozoa, such as the Podo-
coryna, be cut off and the trunk left in contact with
the colony, a fresh head will grow, while the detached
head becomes fixed and forms a new trunk.
In all these phenomena we see a regenerative capacity
of such rapidity and extent that the danger of infection
becomes reduced to a minimum. We may here observe
the regenerative side of inflammatory processes, but not
the phenomena of inflammation itself or at least not the
accumulation of phagocytes at the injured spot.
This accumulation is not however an uncommon
occurrence in the Ccelenterata. Most of these animals,
like the Acrospeda, the Ctenophora and the true Polyps,
are provided with a tolerably well developed mesoderrn,
1 See my article in Arbeiten des zool. Institutes zu Wien,
vol. v. 1883, pp. 143-146.
LECTURE V. 59
and contain in their intercellular substance a number of
amoeboid cells which have all the properties of phago-
cytes.
If we take a large Medusa, known as Rhizostomum
Cuvieri, and introduce some pointed body, as a splinter
of wood or even a pin, into its gelatinous bell, the very
next day it will be perceptible to the naked eye that a
cloudiness has arisen around the foreign body. This,
when microscopically examined, will be found to consist
of numberless amoeboid cells which have collected round
the seat of lesion. The same thing takes place in
another of the Acalepha, Aurelia aurita. If the object
introduced into the bell of the Medusa has been pre-
viously soaked in a colouring matter, such as carmine,
the phagocytes which assemble at the injured spot will
be found filled with coloured granules. The amoeboid
cells which accumulate around the foreign body either
remain isolated or unite to form minute plasmodia.
We therefore see that in these Medusae, which have
no kind of vascular system, the mesodermic phagocytes
are able to traverse a gelatinous substance, which is
sometimes, as in the Rhizostomum, very tough, and to
collect for the purpose of englobing the minute bodies
or of surrounding the larger foreign bodies which have
been introduced.
The analogy of these events with the reactive pheno-
mena in the Sponges is obvious, although there is a
considerable difference between the mesoderms in the
two cases. Whereas in the Sponges, the mesodermic
phagocytes by taking up solid food-particles play an
important part in nutrition, in the Medusae and in all
the Ccelenterata which possess a mesoderm, the func-
tion of nutrition is exclusively confined to the endoderm.
60 INFLAMMATION.
In all the Ccelenterata this consists of a phagocytic
epithelium entirely separate from the mesoderm, at any
rate in the adult condition. Although deprived of their
nutritive function, the mesodermic phagocytes retain
their power of approaching foreign bodies, of englobing or
surrounding them and of digesting some of them. This
power is not only exercised on the foreign bodies which
have penetrated into the Ccelenterata as a result of some
lesion ; it is equally active in the case of the tissues of
these animals themselves. Thus the abortive genera-
tive cells, of which there are a large number in Medusae
kept under artificial conditions, are duly devoured by
the phagocytes, which surround them as with a kind of
follicle. It is apparent that these mesodermic cells
have not lost their primitive properties of intracellular
digestion, and although they have become distinctly
separated from the endoderm, the common origin of the
two layers may be embryologically demonstrated.
The development of the mesodermic amoeboid cells
at the cost of the endoderm which is a fact of very
frequent occurrence in the animal kingdom may be
readily followed in the various representatives of the
Echinodermata, especially in the sea-urchins and the
star-fish. We will take a star-fish common in the Gulf
of Trieste, the Astropecten pentacanthus, as an example.
The segmented ovum becomes converted into an oval
body consisting of a layer of ciliated cells enclosing a
segmentation cavity. A part of these cells becomes in-
vaginated to form the first rudiments of the intestinal
canal and its appendages. The larva soon assumes the
characteristic gastrula stage, and consists of an ectoderm,
or outer layer, and of an endoderm, forming a cul-de-sac
with an opening at the lower end. The space between
LECTURE V. 6l
the two layers represents the general body-cavity and
is filled with a homogeneous and semi-fluid substance,
containing the amoeboid cells of the mesoderm. These
are merely migratory cells which have become budded
off from the endodermic imagination. 1 (Fig. 29.) Al-
most immediately upon their arrival in the body-cavity
they are able to begin their phagocytic duties. Among
the numerous larvae of the Astropecten which float on
the sea, some may be found with their delicate ectoderm
injured by a sharp body which has pierced the general
body-cavity. (Fig. 30.) But as
soon as the larva has been thus
damaged, the mesodermic cells
travel towards the invading object,
and surround it completely by
fusing into minute plasmodia.
(Fig. 31.) The latter, if duly
prepared in a o'5 per cent, solu-
tion of osmic acid and stained
with picrocarmine, are seen to
. . , i r Fig. 29. Formation of phago-
COntam a Certain number Of cytes in the larva of Astropecten.
nuclei, the appearance of which
shows definitely that no proliferation of cells is taking
place. In these larvae, the reaction, which can be
followed step by step owing to their transparency,
consists merely in an accumulation of mesodermic
phagocytes around the foreign body. There can be no
question here of the intervention of any vascular,
muscular, or nervous systems, since in these larvae such
1 Cf. my article in the Zeitschrift f. wissensch. Zoologie^ vol. xlii.
1885. See also the account of the discussion between Selenka and
myself, and the more recent paper by Korschelt in the Zoologische
Jahrbucher, vol. iv. 1889.
62
INFLAMMATION.
systems do not exist. The reaction is effected entirely
by the phagocytes themselves, and is accompanied
neither by proliferation of cells nor by increased flow
of fluid to the part, as shown by the absence of oedema.
The non-occurrence of proliferation is readily explic-
able by the fact that the foreign body on account of its
minuteness has produced but a very slight lesion of the
ectoderm.
In larvae of more highly deve-
loped and complex organisation,
the reaction takes place in the
same manner. I have frequently
seen a marine alga, a species of
Chcetoceros which is provided
with very delicate hairs, settle on
and penetrate into specimens of
Bipinnaria (larvae of Astropecten).
In all these cases, the lesion was
followed by an accumulation
of mesodermic phagocytes to-
ler with the formation of
plasmodia.
In the instances above quoted, the larvae were too
minute to admit of artificial experimentation ; it was
merely a question of watching the effects of lesions
arising under natural conditions. But the reactive
phenomena ensuing on artificial injuries l may be readily
observed in the much larger larvae, the Bipinnaria
asterigera, which likewise represent a stage in the
development of the star-fish. If a delicate glass tube, a
rose-thorn, or a spine of a sea-urchin be introduced into
1 See my article in the Arbeiten des zoolog. Institutes zu
Wien, 1883, vol. v. p. 141.
Fig. 30. Gastrula with a foreign
substance in its body-cavity.
LECTURE V. 63
one of these larvae, the amoeboid cells of the mesoderm
collect around the foreign body in large masses easily
visible with the naked eye. All the minute particles
adherent to the object introduced, or the granules of
carmine or indigo, if the object has been previously
immersed in these substances, are eagerly devoured by
the mesodermic phagocytes. (Fig. 32.)
If instead of these sharp solid bodies, a drop of blood
be introduced into a Bipinnaria, it will be at once
Fig. 31. The oreign body of Fig. 30 surrounded by a plasmodium of
the larva (highly magnified).
surrounded by the mesodermic cells, which will collect
around the masses of blood corpuscles to form true
plasmodia, that is to say, multinuclear protoplasmic
masses arising from the complete fusion of the
phagocytes (Fig. 33). The changes undergone by the
red corpuscles of the blood within the mesodermic cells
of the larva correspond exactly with the phenomena of
intracellular digestion, and may be observed in the same
way with the fat-globules of milk.
Bacteria if introduced into the Bipinnaria are likewise
enclosed by the mesodermic phagocytes. The great
64 INFLAMMATION.
transparency of the larvae of Echinoderms enables the
observer to ascertain the fact that the bacteria are still
mobile, and that they have therefore been devoured
alive.
In spite of the differences which distinguish the
Sponges, the Ccelenterata possessing a mesoderm, and
the Echinoderms from each other, they are all essentially
similar in their phenomena of reaction. In the Sponges
we have a mesoderm with a plentiful supply of mobile
cells which play an important part in the nutrition of
these animals. The food,
after entering the body,
invariably reaches the
mesoderm, which is in-
timately connected with
^ e en doderm. In the
Acale P ha (Medusae) and
the other Ccelenterata in
Fig. 3 2.- r coiiection of phagocytes round a which this layer is present,
splinter. Bipinnaria asterigera. . ,. ,
the mesoderm is directly
connected with the endoderm only while it is being de-
veloped. When its development is complete, the meso-
derm becomes definitely separated from the endoderm and
has nothing further to do with the function of nutrition,
which is relegated entirely to the endodermic phago-
cytes. In the larvae of Echinoderms the two layers are
equally distinct from each other ; the mesoderm is like-
wise excluded from the office of nutrition, while the
endoderm which is the sole organ of nutrition, has no
power of intracellular digestion. In this animal, digestion
is performed by means of ferments secreted by the
endodermic cells, and poured into the intestinal cavity.
Now although these animals differ thus in organisa-
LECTURE V. 65
tion from one another, their mesodermic cells are alike
in that they move towards foreign bodies, englobe and,
when possible, digest them. In all of them the various
lesions produced by such foreign bodies provoke an
accumulation of mesodermic phagocytes, with or with-
out formation of plasmodia or giant cells.
Moreover these animals all resemble each other in the
fact that the phagocytes of their mesoderm are repre-
Fig. 33. Plasmodium formed by the phagocytes of the Bipinnaria,
sented by branched connective tissue corpuscles
embedded in a semi-fluid or gelatinous intercellular
substance. In all the cases that we have considered,
there was neither blood nor plasma, blood-corpuscles
nor blood-vessels. These structures are not found
either in the Sponges or in the Crelenterata, and are
only present in the Echinoderms at a later period of
development than that of the specimens which served for
these pathological investigations.
If now we turn our attention to the varied group of
the Vermes, we shall at once meet with reactive phe-
F
66 INFLAMMATION.
nomena similar in character to those described above^
As a representative of the lower orders of worms, we
will take the transparent Turbellarium, Mesostomum
Ehrenbergi. If we injure any part of its body, the
mesodermic phagocytes will, after a certain time has
elapsed, assemble round the lesion. The number of
granules they contain gives them a great resemblance
to the epithelial cells of the intestine which, in the
Turbellaria, are likewise true phagocytes. The meso-
dermic cells are amoeboid elements embedded in a
gelatinous intercellular substance, forming a mucoid
connective tissue like the mesoderm of the Sponges,
Medusae and Echinoderms.
In the higher Worms the mesodermic phagocytes are
represented by the peritoneal endothelium or by cells
suspended in the perivisceral fluid. These two varieties
of cells have the same marked phagocytic properties,
and this functional analogy may explain the fact that in
closely allied species, the perivisceral cells are some-
times highly developed while at other times they are
completely absent. These mesodermic elements be-
sides fulfilling their phagocytic duties, act as respira-
tory and excretory organs. 1
If a splinter be introduced into the periviscral
cavity of an Annelid, such as the TerebelZa, it will be
soon covered with a thick layer of these 'lymphatic'
cells, the phagocytic properties of which are shown by
the readiness with which they take up the minute
grains (of colouring matter or otherwise) attached to
the splinter. This is the more interesting, since the
1 See Grobben, " Die Pericardialdriise der chaetopoden Anne-
liden." Sitxungsberichte der k. Akad, d. Wissensch., Wien, vol.
xcvii., 1888.
LECTURE V.
67
majority of the Annelida are endowed with a highly
developed and completely closed vascular system. The
reaction to foreign bodies is however confined to the
mesodermic phagocytes, the vessels taking no part
whatever in the process, as may be readily seen owing
to the bright colour of the blood in them.
The same phenomena occur in the Annelida which
Fig. 34. A larva of Gordius encysted and surrounded by a
plasmodium in a specimen of Nais.
have a well-developed vascular system, but no peri-
visceral leucocytes. If a certain number of Nats
proboscidea be examined, some of the individuals will be
found to be infected with larvae of Gordius, which after
entering the general body cavity, excite phagocytic re-
action of the peritoneal cells alone. The latter send out
protoplasmic processes and form minute plasmodia
around the larvae, which protect themselves by secreting
a chitinous covering and enclosing themselves in a sort
of cyst. Although these cysts with their surrounding
F 2
68 INFLAMMATION.
plasmodia are sometimes found in close proximity to the
vessels, the latter do not react in any way to the presence
of the parasite. (Fig. 34.) If any exsudation from the
vessels took place into the interior of the plasmodium,
it would be at once evident, since the blood-plasma is
yellow while the perivisceral fluid is perfectly colour-
less.
As it is manifestly impossible to experiment upon the
minute Nats proboscidea, the phagocytic properties of
the peritoneal cells of this Annelid must be studied
in specimens infected by a microsporidium belonging
to the microbes of ' pebrine.' These parasites excite a
similar reaction on the part of the peritoneal endothe-
lium, the minute spores being taken up by the
phagocytes which have become separated from the
peritoneum. Occasionally these spores may be seen
surrounded by vacuoles as in the most typical cases of
intracellular digestion.
The larger Annelida may likewise be employed for
these researches. Valuable information may be ob-
tained from the study of the common earth-worms,
which are frequently invaded by parasites. Among
these the most frequent and also the best known are the
Gregarinae belonging to the genus Monocystis, which
attack the male organs. When once inside the latter,
these mobile Protozoa have to encounter a large num-
ber of amoeboid cells, which are among the most active
of phagocytes. They are provided with slender and
membranous protoplasmic processes (Fig. 35, A), and
devour with the greatest eagerness all foreign bodies
which come in their way. Even when examined in the
aqueous humour of the rabbit or other inert fluid,
these cells give evidence of their phagocytic activity by
LECTURE V.
6 9
englobing grains of colouring matter or any other
minute bodies that may have been added to the pre-
paration. If they meet with a larger object, such as a
thread of cotton, they will collect in groups and finally
surround it with their protoplasm. (Fig. 35, B.) Now
these same cells react to the parasites which have
penetrated the earth-worms.
While the Gregarinae are in
their condition of activity,
they repulse the phagocytes
by the violence of their move-
ments so that the latter are
rarely able to fix themselves
on the parasite. But as soon
as it attains the quiescent
state, the phagocytes adhere
to its surface, frequently
collecting to form a dense
mass around it. The Gre-
garina evidently objects to
this living covering and seeks
to defend itself by the secre-
tion of a cystic membrane.
(Plate II., Fig. i.) Thus
protected, it begins to produce spores by dividing
into a large number of increasingly smaller oval bodies,
and gives rise to the pseudo-navicellae which have
been so often described. The mass of surrounding
phagocytes continues however to act upon the parasite
and frequently succeeds in injuring and even in killing
it. The encysted Gregarina goes on defending itself by
means of chitinous secretions which assume exaggerated
proportions and become fringed with irregular processes,
Fig. 35. A. Phagocyte of Luntbricus.
B. Collection of phagocytes of Lum-
bricus round a foreign body.
70 INFLAMMATION.
presenting a strikingly abnormal appearance. (Plate
I., Fig. 2.) Finally the whole interior of the en-
cysted Gregarina becomes highly refracting, and the
parasite dies, leaving the phagocytes the masters of the
day. (Plate I., Fig. 3.) The phagocytes themselves
undergo marked alteration around the parasite and,
losing their power of movement, are converted into flat
closely apposed cells. (Plate II., Fig. I, 2.) Sometimes
the capsule thus formed, which has the same structure
as connective tissue, remains very thin ; but it usually
becomes thickened by the further addition of fresh
Fig. 36. A live Rhabditis surrounded by a mass of the phagocytes of Lumbricus.
layers of cells. Among these may be seen some which
are deeply pigmented with a brown colour. During
the whole course of the struggle between the parasite
and the phagocytes of Lumbricus, the blood-vessels
although highly developed remain quite passive, that is
to say, there is no visible change in their volume, nor
any exsudation of the reddish plasma.
The spermathecae of Lumbricus as well as its
general body cavity, may also be invaded by Nema-
toda belonging to the genus Rhabditis. The latter,
in spite of their size and mobility and the toughness
of their cuticle, have likewise to do battle with the
LECTURE V. 71
numerous phagocytes of the earth-worm. These cells
surround the Nematode forming a thick capsule round
it like that which is produced round the Gregarina.
Microscopical examination of the Rhabditis in this
condition proves directly that the phagocytes have
enclosed it while still alive, as it may be seen moving
in the midst of the mass of cells. (Fig. 36.) The
worm, thus confined in its movements, secretes layers
which form, not a true cyst, but a supplementary cuticle
which frequently becomes of extraordinary thickness.
(Plate I., Fig. 4.) This abundant secretion evidently]
Fig. 37- A Rhabditis without its phagocytic covering in order to
show the cuticular processes.
exhausts the parasite, for it loses the granules of fat
with which it was at first filled and becomes quite
transparent. (Plate I., Fig. 5.) The chitinous layers,
as they become thicker and thicker, finally form irre-
gular processes, which give a strange abnormal appear-
ance to the Rhabditis. (Fig. 37. Plate I., Fig. 4.) In
isolating the collections of phagocytes enclosing the
worm with its thickened cuticle, it often happens that
the Nematode manages to escape, leaving its cuticle in
the mass of cells. On the other hand, if the contents
of the male organs of Lumbricus be examined, there
will frequently be found situated in the midst of
phagocytic capsules, highly refracting bodies, which
may readily be identified as the shapeless cuticular
72 INFLAMMATION.
layers and the remains of the Nematode buried in the
products of its own secretion.
Here then we have an example of a struggle be-
tween two members belonging to the same group of
the animal kingdom. The Nematode worm protects
itself by means of cutaneous secretions ; the earth-
worm fights by means of an army of mobile cells, en-
dowed with phagocytic properties. It is evident that
the latter greatly embarrass the parasite by surround-
ing it with their solid masses, although we do not yet
know the exact nature of the harmful influence exer-
cised by the phagocytes on the intruder. They may
act by preventing the inflow of nutritive material or
of oxygen, or by means of an injurious secretion.
These delicate points can only be decided by the
most minute research, and must await a perfection of
our methods for their final answer.
At present we must be satisfied with the statement
that Lumbricus, like the Annelida generally, reacts
to various infective agents by means of the phago-
cytes of their perivisceral fluid without any intervention
on the part of the blood or of the highly developed
blood-vessels. This reaction takes place in the same
way against Gregarinse as against Nematoda. In
speaking of the latter, I must particularly mention
the fact that they are devoid of migratory cells. The
pjhagocytic system of the Nematoda is probably re-
duced tcT the muscular phagocytes, which are~1ormed
in a very_ cur i us manner." TKese animals protect
themselves by the secretion of tough membranous
cuticles, resembling in this feature the plants, the cells
of which are likewise^rote^teoV-^rtfilcirresistant mem-
branes. This analogy is borne out by the fact that
LECTURE V. 73
these animals, like plants, are very frequently attacked
by parasitic fungi, which are enabled to penetrate the
cuticle by their great power of growth and also by
the secretion of ferments which can dissolve the most
impervious substances, such as cellulose.
Among the infectiousdiseases of the Nematoda, we
must name one which is produced by the parasitism
of one of the Mucorineae (Mucor helminthophorus, De
Bary), which invades the intestine and the genital
organs of the Ascaris of the cat (A. mystax}? as well
as the frequent infections of the free Nematoda by
several other members of the class of fungi. The
most remarkable is certainly the Arthrobothrys oligos-
pora Fres., because, according to Zopf, 2 this mould
catches the Anguillulides with hooks, and afterwards
penetrates their bodies with its filaments. Once inside,
the fungus grows freely in the body cavity and causes
complete fatty degeneration followed by death of the
animal. There is finally nothing left of the Nematode
but its cuticle and the chitinous covering of the male
genital organs.
Besides these epidemics, occasioned by the higher
fungi, the Nematoda are liable to invasion by Chytri-
diaceae and other inferior organisms, allied to those
which infect the nucleus and nucleolus of the Para-
maecia. 3
This survey of the pathological phenomena in
Ccelenterata, Echinoderms and Worms, has shown
1 Zeitschrift f. wissensch. ZooL, 1862, vol. ii. p. 135.
2 Nova actuAcad. Leopold, vol. xlvii. p. 167; and "Pilze," 1890,
p. 240.
3 See Biitschli, " Studien \iber die ersten Entwickelungsvorgange
der Eizelle.' J Frankfurt, 1876, p. 360.
74 INFLAMMATION.
that some of these animals react chiefly by the rapid
and active regeneration of the injured parts, while
others protect themselves by secretions of chitinous
layers. These two methods, however, only hold in
certain cases, whereas the usual mode of reaction, to
which there are but few exceptions, is by means of
amoeboid and mobile cells which accumulate around
the injurious body and either surround it entirely or
englobe it. The reaction is effected through the sensi-
bility of the phagocytic cells themselves, and is in no
way influenced by the nervous or vascular system. In
all the above-mentioned cases, the phagocytes were
mobile connective tissue cells, or cells of the peri-
visceral cavity. So far we have not discovered any
instance of phagocytic action on the part of the blood-
corpuscles. It is true that our observations have been
confined to animals which have no formed elements
in the blood. Annelida with white corpuscles in
their blood do not often occur, and even in cases
where these are present, their number is small, and
certainly less than the amount found in the perivisceral
cavity.
LECTURE VI.
Arthropoda, Molhtsca, and Tunicata Their vascular system
Their Phagocytes Spleen of the Gasteropoda Inflammatory
reaction Diapedesis in intact Ascidians Introduction of
bacteria into the body of Ascidians and Crustaceans Infec-
tious disease of sandhopper (Talitrus) Diseases of Daphnia
Introduction of bacteria into insects Epidemics among
insects.
A LARGE number of the invertebrata possess blood-
corpuscles in the form of colourless cells which float in
the blood-plasma. The circulation of this fluid is
effected by the movements of the heart, which is always
present. In these animals Arthropoda, Mollusca, and
Tunicata the vascular cavity is identical with the
general body cavity. In the lower representatives of
these types (we disregard certain groups without a trace
of a vascular system, such as many of the Copepoda,
Ostracoda and others) the only vascular organ present
is the heart, in the form of a simple sac or tube open at
its extremities to expel the blood and provided with
lateral apertures for the entry of this liquid. To this
central organ are soon added one or several principal
arteries which open into a system of lacunae, in which
the blood circulates before going back to the heart.
In the invertebrata rather higher in the scale, especially
in the Mollusca, we find also a venous system, which
-is sometimes, as in the Cephalopoda, very highly
76 INFLAMMATION.
developed. But in all cases without exception, even
when a large number of vascular ramifications are
formed, there is a network of lacunae between the
arterial and venous systems. These lacunae are filled
with blood and are remains of the general body
cavity.
The blood- corpuscles with a few rare exceptions
are represented by colourless cells, possessing one or
rarely two nuclei and a protoplasmic body capable of
amoeboid movements. In many invertebrates there is
only one variety of mobile blood-corpuscles, containing
a few sparse granules, whereas in certain others, such
as many insects and molluscs, two varieties occur
granular leucocytes, with a large number of coarse
granules, and hyaline leucocytes, with few or no
granules. The latter kind is the one which will interest
us most.
The leucocytes of Arthropoda, Mollusca, and
Tunicata are in most cases amoeboid and phagocytic
cells, and differ from the white corpuscles of verte-
brates in having a single round or oval nucleus, which
is not lobed. In the invertebrates now under considera-
tion no multinuclear leucocytes exist, nor do we find
in them a vascular system with complete capillary
blood-vessels.
The leucocytes of the three types just mentioned
manifest pronounced phagocytic functions. It was in
a representative of these invertebrate groups that the
discovery was first made (in 1862) that leucocytes
possessed the power of taking up foreign bodies into
their interior. Haeckel l showed that, after injecting
the mollusc Thethys with indigo, granules of
1 "Die Radiolarien," 1862, p. 104.
LECTURE VI. 77
this colour were to be found within the blood-cor-
puscles. Experiments with several other species led
him to the conclusion that this was a fact capable of
general application a deduction which has since been
confirmed by several observers. It is therefore very
surprising that a recent author, Griesbach, * should
throw doubts on the occurrence of phagocytosis
in the white corpuscles of the acephalous Molluscs.
Since he did not succeed in observing any considerable
taking up of a powder after injecting it mixed with
water, Griesbach concludes that under normal condi-
tions phagocytosis does not occur at all in these
animals. It is very probable that the unsatisfactory
results of this author were due to the fact that he used
too much water to dilute the powder, and so caused the
leucocytes to swell up. If we proceed more carefully,
it is easy to show that in the Molluscs, as in so many
other animals, the leucocytes take up greedily any solid
bodies with which they happen to be in contact. The
transparent Molluscs such as Philliroe, which admit of
direct examination under the microscope in the living
condition, form very convenient objects for these
researches.
In some gasteropod Molluscs, we find in addition to
the white corpuscles, a special variety of phagocytes
which form a kind of spleen in these invertebrata.
This important fact has been recently discovered by
A. Kowalewski, 2 who has shown that solid bodies,
injected into the blood of the Pleurobranchsea and
several other species (Philina, Gasteropteron, Doris),
1 Archiv f. mikroskop. Anatomic, vol. xxxvii. p. 86.
2 Memoires de la Societe des Naturalistes de la Nonvelle Russie,
vol. xv., 1890 (in Russian).
78 INFLAMMATION.
accumulate in a certain organ which was first described
by de Lacaze-Duthiers under the name of " glande
indeterminee." The cells of this 'spleen/ as
Kowalewski has established, deovur and digest a large
number of foreign bodies such as blood corpuscles, yolk
granules, and milk corpuscles.
The phagocytes, so universally present in the inverte-
brata which form the subject of the present lecture,
react to all sorts of lesions, whether these are artificially
or accidentally produced in these animals. We often
eome across transparent Crustacea, such as Daphnia or
Branchipus, with brown spots on their sides, due to
bites inflicted by other
individuals. Under-
neath these eschars we
generally find amass of
leucocytes, which re-
main heaped together
Fig. 38,-Inflamed cauda appendage of Argulus. at the mjliredspOt Until
the wound is com-
pletely cured. 1 If we carefully inflict a small injury on
one of these animals and observe it under the micro-
scope, we see the leucocytes making towards the
affected spot, where they take up their abode. A con-
venient object for experiments of this sort is furnished
by the caudal appendages of Argulus, in which the leuco-
cytes collect directly after the production of the artificial
lesion (Fig. 38).
We may also introduce fairly large foreign bodies,
such as wooden splinters, into the larvae of various
Coleoptera (cockchafer, Oryctes and others), into
1 The extremely rapid regeneration of the epidermis in the
Arthropoda causes their wounds to heal very quickly.
LECTURE VI. 79
molluscs such as Thethys or Phylliroe or into Ascidians. 1
In all these cases a great number of leucocytes may be
seen to collect around the foreign body, devouring any
little fragments or granules as, for example, carmine,
that may have been introduced with it.
In all cases then a phagocytic reaction is produced
by the introduction of a foreign body, the leucocytes
either forming a capsule round the intruder, or infil-
trating all the surrounding tissues. In this exsudative
and inflammatory reaction, which is often accompanied
with the formation of giant cells, diapedesis can play
no part, for the simple reason that in the Arthropoda
and Mollusca there is no closed vascular system
properly speaking, the blood cavities being merely part
of the general body cavity.
In the invertebrata only one single example of dia-
pedesis occurs ; but this is of so interesting a character
that it deserves a detailed description. The Ascidians
are covered with a mantle or tunic which is situated
outside the epidermis. This tunic, which is composed
of cellulose and is often very thick, contains a large
number of amoeboid cells with mobile processes. Since
it is situated outside the epidermis, it was generally
regarded as a cutaneous secretion containing cells of
ectodermic origin. Later researches by Kowalewski 3
have shown that this view was unfounded, and that the
cells in the tunic of Ascidians are in reality nothing
else than emigrated leucocytes which have wandered
through the epidermis. These cells of mesodermic
origin are very active phagocytes, and are capable of
devouring all sorts of solid bodies, including organs
1 See fatArbeiten des zoolog. Inst.zu Wien,\o\. v., 1883, p. 153.
- Loc. cit.
80 INFLAMMATION.
that are undergoing atrophy. Insertion of splinters
into the tunic of Ascidians provokes an accumulation
of these phagocytes, so that a sort of infiltration of the
tunic is produced.
We have here an example of diapedesis taking place
under normal conditions through the epidermis, quite
independently of any inflammation; though this latter
process is also carried out by the aid of the same
phagocytes, which collect around the offending particles.
Lubarsch ' has confirmed the observation that the
mobile cells in the tunic of Ascidians congregate around
foreign bodies that have been introduced by a puncture.
He was not so successful in his experiments on the
inoculation of various Ascidians with the bacilli of
anthrax. The bacteria which had been introduced
into the tunic were only partially taken up by the
phagocytes, while those which escaped this fate never-
theless showed marked signs of degeneration. As
Lubarsch did not investigate the direct influence of the
fluid portions of the tunic on the bacteria, we cannot
form any definite conclusion from his none too
numerous experiments. In considering them we must
not lose sight of the facts that they were carried out in
the month of March, when the low temperature might
have had an injurious influence on the leucocytes, and
that the tunic of Ascidians cannot afford a very favour-
able soil for the growth of bacteria and the production
of their toxines. 2 Lubarsch 3 has also made some
1 " Untersuchungen liber die Ursachen der angeborenen und
erworbenen Immunitat." Berlin, 1891^.75.
2 I must here mention the fact that the tunics of compound
Ascidians such as Botryllus, often show phagocytes filled with
various bacteria, even when the animals are examined immediately
after being taken from the sea.
3 Loc. tit., p. 77.
LECTURE VI. 8 1
similar experiments on " marine Crustacea " without
any better result. The experiments are recorded very
shortly, so that it is impossible to criticise them. We
have a number of facts, however, which prove in
the clearest manner the pronounced phagocytic
properties possessed by the leucocytes of different
Crustacea. By the introduction of a parasitic fungus
(allied to Oi'dium) into the body cavities of sand-
hoppers (Talitrus), Hermann and Canu ' have suc-
ceeded in producing a disease, which is almost always
fatal to these Crustacea. The development of the
parasite excites reactive changes on the part of the
organism, as shown by a pronounced phagocytosis of
the leucocytes. The authors describe these changes as
follows : " On the seventh day, the blood, which up to
this time has been quite clear, becomes appreciably
opalescent, and the disorder becomes more marked on
the eighth and ninth days, as the parasites increase
in number. This is also the period at which the
most active phagocytosis is observed ; if the blood,
fixed by osmic acid vapour and stained with picrocar-
mine, be examined under a high power, the microbes
are seen to be enclosed in the corpuscles, the number
in each corpuscle varying from one to twenty. In the
protoplasm they are seen to be in various stages of
digestion ; they become paler and less highly refracting
at the same time that they increase in size, chiefly in
consequence of a swelling up of their enveloping mem-
brane. Finally the place they occupied in the cell is
marked only by a sort oi colourless vacuole which
preserves for a considerable time the elongated shape
of the parasite." Besides the blood-corpuscles, the cells
1 Coniptcs rendus de la Societe, de Biologic, 1891, p. 646.
G
82
INFLAMMATION.
surrounding the arteries also take part in devouring
the parasitic fungi, thus acting as phagocytes, although
they are not able to digest their prey. In the end,
therefore, the parasite gains the upper hand and brings
about the death of the sandhopper.
Luminous bacteria also, as Giard and Billet have
shown, live as parasites on the same species of
Amphipoda.
Fig- 39- Daphnia infested by Monosporae.
A large number of the Crustacea are subject to
various infectious diseases which form an interesting
subject of study from the pathological point of view,
and especially on account of the light they throw on
the problem of inflammation. The Daphniae afford
especially convenient objects for these researches, in
consequence of their transparency and minute size, and
the frequency and variety of their diseases. Among
the latter we find infectious maladies produced by
bacteria, Sporozoa or Saprolegniae. The most interest-
ing of these, however, is certainly that provoked by the
presence of a fungus multiplying by budding, Monospora
LECTURE VI.
bicuspidata? This is a sort of yeast, which is found in
abundance infesting Daphnia magna in Paris (in the
reptile tank of the Jardin des Plantes) and its
environs.
Among the numerous individuals of this Crustacea,
we come across specimens distinguished by their milk-
white colour. On examining these under the micro-
scope, we see that their body-cavity is almost entirely
filled with small needle-shaped bodies, either floating
freely or adhering to the walls of
the heart (Fig. 39). A close in-
spection shows at once that we
have here very long spores, en-
closed in a capsule (Fig. 40, 5).
By the side of these mature
spores, elongated cells and oval
conidia are seen multiplying by
budding, exactly in the same
manner as the yeasts. (Fig. 40,
1-4.)
A Daphnia, once invaded by
these parasites, always dies,
and its body is found filled with ripe spores. Other
Daphniee, feeding on every sort of detritus which they
find at the bottom of the water, devour the needle-
shaped spores, and thus infect themselves through their
alimentary canal. In the intestine the spores lose their
capsule, and penetrate the wall so as to lie partly or
entirely in the general body-cavity of the Crustacea.
Directly the spore, however, appears outside the
intestinal wall, it is attacked by leucocytes, which are
Fig. 40. Monospora in various
stages.
i. Young conidium. 2, 3. Bud-
ding conidia. 4. Elongated
conidium. 5. Spore.
1 VircJww's Archiv., vol. xcvi. p. 177.
G 2
8 4
INFLAMMATION.
carried to the spot by the blood-stream. These cells
fix themselves on the spore, forming around it a collec-
tion of cells, which often fuse together into a plas-
modium. Under this influence the spore undergoes a
series of remarkable changes. On being enclosed in
the leucocytes the spore first loses its regular contour,
becomes sinuous, and finally breaks up into a mass of
brownish granules in which it would be impossible to re-
cognise the degenerated spores if we had not studied the
mode of their formation. (Fig. 41, 1-4.) The proof
that this degeneration is
really due to a phagocy-
tic influence is furnished
by those cases in which
only half a spore is sur-
rounded by leucocytes,
the other half being
embedded in the in-
testinal wall, or even
projecting outside the
skin of the animal (Fig. 41, 5). In these cases it is only
the half surrounded by the phagocytes that undergoes
the changes just described, while the other half, which
is not exposed to the influence of these cells, remains
perfectly normal.
When a large number of spores
penetrates the body cavity of the
Daphnia at one time, the leuco-
cytes congregate round them, in
a manner resembling an infiltration or a cellular exsuda-
tion (Fig. 43, a). It is in fact a sequence of events
exactly similar to that which we have seen produced by
a traumatic lesion.
Fig. 41. Spores of Monospora, surrounded
by the leucocytes of Daphnia.
Fig. 42. Elongated conidium
of Monospora surrounded by
two leucocytes.
LECTURE VI. 85
The phagocytic action of the leucocytes, so evident
and easily studied in the transparent Daphniae, destroys
the spores of the pathogenic microbe and prevents their
development, thus protecting the invaded organism. I
have succeeded several times in isolating infected
Daphniae and keeping them till they were fully restored
Fig. 43. Hind part of a Daphnia.
a. Spores of Monospora surrounded by a mass of leucocytes.
to health, thanks to the destruction of the spores by
their phagocytes. If on the other hand the phago-
cytic action is inadequate, owing to the continued in-
crease in the number of spores swallowed or for any
other reason, the latter begin to germinate and give rise
to budding conidia. Although the parasite in its
vegetative form is also attacked by leucocytes, it
obtains the upper hand and the Daphnia inevitably
86 INFLAMMATION.
succumbs in a short time to its attack. This is owing
to the fact that the conidia multiply too rapidly, and
also secrete some poison which dissolves the leucocytes.
Towards the end of the disease nothing but conidia are
to be seen circulating in the body of the Daphnia, all
the leucocytes having completely disappeared.
There can be no doubt that the whole history of this
disease of the Daphniae may be summed up as a struggle
between two living organisms, the parasitic cells and
the phagocytes. In spite of the extraordinary activity
of the former, it is the Daphnia in most cases which,
under the protection of its phagocytes, gains the upper
hand. We can in this way explain the fact that even
when an epidemic of Monospora is raging in a tank or
aquarium containing Daphniae, the number of these
remains as high as ever. While some individuals die
every day, stricken with the disease, the rest resist its
ravages and multiply, thus filling up the gaps caused by
the deaths.
Very different is the course of the diseases caused
by parasites which do not meet with any resistance on
the part of the leucocytes. To this class belong the
Saprolegnise. The spores of these fungi germinate on
the surface of Daphniae or other Crustacea (as e.g.
Branchipus) and put forth filaments which penetrate
into their body. The filament to effect an entry often
makes use of the little fissures or orifices produced by
various causes, which may be wounds or little canals
pierced by the spores of Monospora. Once inside the
body cavity of the Crustacea, the Saprolegnia continues
its development in the blood with which this is filled,
meeting with no obstacle from any side. The leuco-
cytes manifest great indifference towards the developing
LECTURE VI. 87
mycelium, which in the end dissolves these cells and in-
evitably brings about the death of the infested animal.
If an epidemic of Saprolegniae has once broken out in
an aquarium, we may be quite certain that it will not
stop until it has destroyed the whole of the specimens of
Daphniae or Branchipus present. 1
Several of the diseases of Daphniae, such as those
caused by the bacteria Pasteuria ramosa 2 or Spiro-
bacillus Cienkowskii* or by the Sporozoa (pebrine and
others), meet with but feeble resistance on the part of
the phagocytes. As we should expect under the cir-
cumstances these diseases, when once established, are
never cured, and infallibly end in the death of the
animals attacked.
The feebleness of the phagocytic protection which is
so striking in Crustacea, is probably connected with the
thickness of their cuticular envelopes, which clothe not
only the external surface but also the intestine in these
animals. The chitinous cuticle is very tough and is
quite impermeable to most microbes. Thus we see
that the small Crustacea, such as certain Copepoda,
which are provided with a very hard covering, can get
on perfectly well without phagocytes, and in fact do
not possess any corpuscles at all in their blood.
The insects, so for as concerns inflammation and
resistance to microbic infection, are exactly similar to
the Crustacea. Every kind of lesion produces in them
an accumulation of leucocytes round the injured spot, as
may be easily seen on cauterising the tips of the caudal
1 Branchipus and Artemia are subject to the disease produced
by the Monospora. The pathological phenomena in these cases
need further investigation.
- Annales de rinstitut Pasteur, 1888, p. 165.
* Ibid., 1889, p. 265.
88 INFLAMMATION.
appendages in the larvae of Ephemeridae or other in-
sects.
Balbiani ' has published some very interesting re-
searches on the effects of introducing bacteria into the
bodies of various insects and Arachnidae, and has found
that many saprophytic bacteria are pathogenic and even
fatal for a large number of these Arthropoda. But
while the insects rich in leucocytes, (such as certain
Orthoptera, especially the Gryllidae,)can completely with-
stand the introduction of a great number of bacilli, the
other kinds that are poor in blood and leucocytes (such
as the Lepidoptera, Diptera, and Hymenoptera,) are
extremely susceptible to infection by the saprophytic
fungi. The power of resistance possessed by the in-
sects belonging to the first order " must be ascribed to
the action on the bacilli exercised by two varieties
of cells. The first of these is represented by the
blood-corpuscles which, by means of their pseudo-
podia, seize on the bacilli floating in the blood and
include them in their protoplasm, where they are quickly
destroyed ; the second variety, the elements of the peri-
cardial tissue, consists of large cells with multiple nuclei,
which surround the heart or dorsal vessel in the form of
plates or cords of cells, and are more or less highly
developed in different types. Of all the tissues of the
body, the pericardial tissue is the only one that has the
power of arresting the bacilli carried by the blood, and
of taking them into their protoplasm, where they are
destroyed as rapidly as in the blood-corpuscles." ~
The insects, although so susceptible to infection by
the most widespread and apparently inoffensive of bac-
1 Comptes rendus de f Academic des Sciences, vol. ciii. p. 952.
" Loc, tit., p. 953.
LECTURE VI. 89
teria, are nevertheless very rarely subject to epidemics
of bacterial origin. The cause of this is probably to be
sought in the fact that the bacilli lack the means of
getting through the thick cuticular wall which covers the
skin and lines the intestinal canal and tracheae of insects.
Besides the lethargy of silkworms (' flacherie '), discovered
by Pasteur, 1 in which infection takes place through the
intestine, there are certain other diseases of insect larvae
which are occasioned by bacteria. Thus the larvae of
Anisoplia austriaca in the south of Russia are some-
times invaded by a bacillus which, in its length and
curved shape, recalls the appearance of the anthrax
bacillus. At the beginning of the disease the affected
larvae are not to be distinguished from normal indi-
viduals, and it is only after the complete invasion of the
blood that they show signs of illness and shortly after-
wards die.
These diseases, however, are not nearly so frequent
as those caused by the higher fungi or by the Sporozoa,
which are much better adapted than the bacteria for
penetrating the chitinous coverings of insects. In the
strength of their growth the former possess a powerful
means whereby to penetrate the cuticular walls, while
the Sporozoa have an amoeboid stage during which, by
virtue of their mobility, they are able to effect an entry
at even the most protected spots.
An observation of de Bary 2 on the conidia of
Cordiceps viilitaris that had been englobed by the
leucocytes of caterpillars, leads us to conclude that the
conidia of fungi which multiply in the insect blood,
1 " litudes sur la maladie des vers a sole," 1870, vol. i.
2 " Vergleichende Morphologic und Biologic der Pilze," 1884,
P- 399-
90
INFLAMMATION.
sometimes meet with a certain amount of resistance on
the part of the phagocytes. But in the majority of
cases that have been examined to decide this point, the
mycelial filaments and conidia develop in the blood
without check or hindrance. I can vouch for this
especially in the case where Cleonus punctiventris, as
larva, chrysalis or perfect beetle, is invaded by the
Isaria destructor. The oval green spore germinates on
the surface of the body, and gives rise to a small fila-
ment. This latter experiences great difficulty in pene-
Fig. 44. A leucocyte of Cleonus, showing two phases of movement.
The conidia of Isaria lie by its side, and are not englobed.
trating the cuticle, which becomes brown round the
puncture made by the parasite. But as soon as this
obstacle is overcome, the filament gains access to the
body cavity where, bathed in the blood, it grows freely.
The leucocytes at times approach the filament or de-
tached conidia, but do not englobe a single one of these
parasites (Fig. 44, 45). The latter therefore speedily
invade the whole animal and transform it into the hard
mass, so characteristic of the dead bodies of the
insects that have perished from one of the varieties of
' muscardine/ These pests, which have but one ob-
stacle, the tough cuticular wall, to overcome, are often the
LECTURE VI. QI
cause of devastating epidemics among insects. We
may call to mind the losses that were formerly occa-
sioned by the ' muscardine ' of the silkworm. The
epidemic disease caused by the Isaria destructor affects
several species of beetles, especially Cleonus punctiven-
tris. It often happens that more than half these insects
Fig. 45. Freeconidia of Isaria in very close promimity to some leucocytes
of Cleonus.
perish from the attacks of the parasite. These insects
do a large amount of damage to the beetroot, and the
owners of the beet plantations in south-west Russia
make their calculations as to the amount of seed that it
will be necessary to sow, according to the rate of mor-
tality among these insects caused by the " muscardine
verte." In fact it is now a rooted conviction that the
culture of beetroot in the district mentioned would be
92 INFLAMMATION.
impossible were it not for the assistance afforded naturally
by the Isaria destructor.
The diseases of insects occasioned by Sporozoa, (a
well-known example being the pcbrine of silkworms,)
have not yet been studied from the standpoint of the
comparative pathology of inflammation. We are
acquainted with the microsporidium 1 which is the
cause of the pebrine, and with its amoeboid condition
which enables it to attain access to various cells, such
as the young eggs ; but the question whether any con-
test takes place between the parasite and the phago-
cytes has not yet been investigated. In Daphnia, which
is equally subject to the attack of the microsporidia,
the resistance offered by the leucocytes is extremely
feeble, and is only displayed against the spores. The
amoeboid condition of the parasites is developed in the
close neighbourhood of the leucocytes, without exciting
them to take any part whatever in checking the course
of the disease. The microsporidia, which develop so
freely in the body cavity of Daphnia, finally invading
the entire animal, have absolutely no destructive action
on the leucocytes. The latter circulate in the blood,
now and then fastening on to the surface of the
parasites and dropping off again as if they had to do
with some harmless object.
In reviewing this chapter on the reactive phenomena
presented by the invertebrata that are provided with
amoeboid and phagocytic blood-corpuscles, we are
bound to conclude that in all these animals an agglomera-
tion of these cells is produced round any injured spots.
This inflammatory reaction takes place as a result
1 See especially Balbiani, " Legons sur les Sporozoaires," 1884,
p. 150, et seq.
LECTURE VT. 93
of any sort oi traumatism (cauterisation, introduction
of splinters, bites, &c.). It is seen also in the course of
certain infectious diseases, as for example that caused
in Daphnia by the presence of Monospora. In the
case where the phagocytic inflammation occurs to any
considerable extent, the cell accumulation consists
chiefly of the leucocytes which are brought along by
the blood current and are attracted to the injured spot
in virtue of their sensibility. The lacunar circulation
aids the approach of the leucocytes and renders un-
necessary any special arrangements for the passage of
these cells, such as we find in the vertebrata.
As, however, the sensibility (chemiotactic or other-
wise) of the leucocytes very frequently remains negative,
highly favourable conditions then exist for the inroads
of all kinds of parasites. In these cases protection
against infection is chiefly provided for by the thick
chitinous integument with which the animal is invested,
so that we have, in the Arthropoda, a means of defence
analogous to that which we have seen to be possessed by
Nematoda and plants. In this type of invertebrata,
however, members entirely devoid of leucocytes arc
extremely rare, the large majority of the Arthropoda
having a more or less highly organised army of these
defenders.
LECTURE VII.
Vertebrata Amphioxus Embryos of Axolotl Young larvae of
Urodela Comparison with the invertebrata Tadpoles
Diapedesis Migratory cells Fixed cells Phagocytic pro-
perties of leucocytes Do fixed cells also functionate as
phagocytes ? Transformation of leucocytes into fixed connec-
tive tissue cells Fate of the leucocytes that do not undergo
this transformation Evolution of inflammation in the organic
world.
THE last survivor of the lower vertebrates, the A mpJii-
oxus lanceolatus, is curiously distinguished from all its
congeners so far as regards its pathology. It possesses
no blood corpuscles whatever, and is only furnished
with a very small number of amoeboid connective tissue
cells. All attempts therefore to provoke inflammatory
phenomena in it have given only negative results. The
application of nitrate of silver, or an incision does not
excite any visible reaction. This is evidently due to
the fact that Amphioxus is possessed of a very tough
limiting membrane, which serves as an important means
of defence to the animal. In this respect it resembles
the Nematoda and other animals that are protected by
their chitinous cuticles, as well as most plants.
In order to obtain reactive phenomena analogous to
those which take place in most invertebrata possessing
a mesoderm, we might turn our attention to the class of
LECTURE VII. 95
fishes, where we find inflammatory processes similar to
those that are known from the study of the higher
animals. Since, however, the fishes are ill-adapted for
investigation in the living condition, it is better to pass
at once to the amphibia. In this class the larval stages
serve as classical objects for researches of this nature,
the caudal fins of the larvae of Urodela (Tritons and
Axolotls) and the tadpoles of Batrachians offering by
far the best material for the investigation of inflammation
in vertebrata.
We will first consider the Urodela, which form the
lowest group of the amphibia.
In the embryo of Axolotl 1 the rudiment of the fin is
completely devoid of blood-vessels and lymphatics. In
addition to the epidermis, it is composed of a layer of
mesodermic cells which are divided at an early stage
into two varieties : fixed cells with processes which
ramify like a stag's horn, and amoeboid cells with large
mobile processes having few or no branches. Although
the fixed cells form a majority of the elements of this
connective tissue, the migratory cells occur in fairly
large numbers. (Fig. 46.)
The embryos of Axolotl, if freed from the egg mem-
branes at the tenth or twelfth day of development,
live readily in the aquarium, and can be used for experi-
ments on inflammation. If in one of these embryos
(previously curarised), we touch the edge of the fin with
a fragment of silver nitrate, and at once wash this off"
with a stream of saline solution, we obtain a small
limited burn ; or we may produce a small lesion of the
fin, by introducing a needle charged with powered car-
1 I have always used the white variety, since these ar more
convenient for researches on inflammation.
9 6
INFLAMMATION.
mine or indigo. By either method we kill a certain
number of cells, and lay bare a part of the tissue of the
fin, which takes up a certain quantity of water, so that
the adjoining cells, especially the stellate cells, become
altered and lose to a great extent their highly refract-
ing appearance and their vacuoles. A short time after
Fig. 46. Connective tissue in the fin of an Axoloil embryo,
a. Amoeboid cells.
the operation we may see a certain number of migratory
cells making towards the injured spot, while the epider-
mis folds over and covers the wound. The next day
some of the amoeboid cells of the connective tissue
accumulate round the injured spot and englobe either
the coloured granules lying in the wound or the
debris of the destroyed cells (Fig. 47). In this collec-
tion of cells some are to be found in the process of
LECTURE VII.
97
karyokinetic division. The number of mitotic figures
however is too small to permit of our ascribing many of
the cells assembled at the seat of injury to the division
of pre-existing ones. Moreover this hypothesis is un-
necessary since direct observation shows clearly that
it is the mobile cells which accumulate round the lesion.
The stellate connective tissue cells, which can be
watched from day to day in the same animal, are entirely
Ifaus*
YJlicktt .re
Fig. 47. Inflamed spot in an Axolotl embryo.
passive in their behaviour. The karyokinetic pheno-
mena seen in some of them, are not more pronounced
than those occurring in other parts of the fin. The
blood-vessels do not take any part in the changes con-
sequent on the injury. Although they form large
trunks in the tail, they are quite absent in the fin or are
present only as small tubes which have no connection
with the rest of the circulation.
Here then in the embryo of a vertebrate animal we
have the phenomena of reaction carried out by the
mobile cells of the connective tissue alone, without any
H
98 INFLAMMATION.
intervention on the part of the blood-vessels or of the
white blood-corpuscles. There can be no doubt that
this process is analogous with the reaction which, as
we have seen, is the result of lesions in the Medusae and
the larvae of Echinoderms ; in both cases there is an
accumulation of the phagocytes of the connective tissue
round the seat of injury.
Phenomena of the same nature may be also observed
in the young larvae of Axolotl, which have blood-vessels
in their fins, as well in the larvae of Tritons, which are
also provided with capillary blood-vessels. Emigration
scarcely or never occurs, owing to the early vessels of
these larvae being too minute to allow of the passage
through their walls of the large leucocytes, impeded in
their movements as they are by the voluminous red
corpuscles. Hence arises a remarkable series of pheno-
mena. The vessels of the fin in the vicinity of the
injured spot remain passive, and do not even undergo
any appreciable dilatation, while the migratory cells
of the connective tissue travel towards the seat of the
lesion.
As the occurrence of an inflammatory reaction in
vertebra without the intervention of vessels is a fact of
great importance, I will devote further attention to the
same phenomenon in the young larva of Triton tczniatus.
The edge of the caudal fin of one of these larvae was
touched with a very small crystal of nitrate of silver,
and washed immediately with salt solution and then
with pure water, so that the lesion was confined to a
small group of epidermic and connective tissue cells.
The branched cells of the connective tissue that
were nearest to the injured spot, became less highly
refracting and absorbed fluid, so that their protoplasm
LECTURE VII.
99
became vacuolated, while at the same time their processes
were drawn in and shortened. (Fig. 48.) Two amoeboid
cells in the adjoining parts of the connective tissue began
to move towards the seat of injury. In a subjacent
capillary loop the circulation was completely arrested.
Three-quarters of an hour after the application of the
Fig. 48. Part of the caudal fin of a Triton embryo, a quarter of an hour
after the application of nitrate of silver.
1-5. Fixed connective tissue cells. a, 6. Migratory cells.
nitrate, the amoeboid cells in the tissues nearest the lesion
had increased somewhat in number, and all were making
their way towards the cauterised spot. (Fig. 49.) Three
hours after the beginning of the experiment the branched
cells still retained their relative positions, but had re-
gained their normal refractive power, and hardly showed
any change in the shape of even their finest ramifica-
tions. The migratory cells continued to increase in
H 2
IOO
INFLAMMATION.
number, but solely at the expense of the neighbouring
connective tissue, since in all the vessels the circulation
appeared to have resumed its normal course, and no
trace of diapedesis was to be seen. Two hours later
i.e. five hours after the cauterisation), the eschar formed
by the necrosed epidermis came away, disclosing a new
Fig. 49. The same specimen, three-quarters of an hour after cauterisation.
1-5. Fixed cells. a, 6, c. Migratory cells.
epidermic layer, beneath which lay a mass of migratory
cells. (Fig. 50.) Other mobile cells were proceeding
in the same direction, whereas the fixed connective
tissue cells retained their former positions. The cir-
culation was restored in the neighbouring vascular loop
and yet, as throughout the experiment, there was abso-
lutely no diapedesis. On the following day the injured
part was completely restored to a normal condition.
LECTURE VII. 101
The stellate cells (Fig. 51) had resumed their wonted
Fig. 50. The same spot five hours after cauterisati
1-5. The same fixed cells as in Fig. 47 and 48.
appearance and again presented their characteristic
antler-like processes. The migratory cells, of which a
certain number remained accumulated under the re-
102
INFLAMMATION.
generated epidermis, were as sparsely scattered in the
connective tissue as in a normal animal.
On seeing how quickly this restitution took place, I
again cauterised the same spot, twenty-four hours after
the first experiment, with nitrate of silver but more
Arrnls so.
Fig. 51. The same spot 24 hours after the cauterisation.
1-5. The same fixed cells as in Fig. 47-49.
severely than on the former occasion. Directly after-
wards the stationary cells underwent the same changes
as before ; they became larger, vacuolated, and less
refractive, while their processes were considerably
shortened. The circulation in several adjoining capil-
lary loops was arrested ; but, in spite of the severity of
the lesion, diapedesis only occurred to an insignificant
degree. During five hours of observation, I saw only a
LECTURE VII. IO3
single leucocyte pass through the capillary walls. This
fact however did not hinder the accumulation of
migratory cells, a certain number of which made their
way towards the seat of injury.
These observations, which have been repeated for
several years with uniform results, demonstrate clearly
the possibility of an inflammatory reaction in vertebrata,
without the intervention of the vascular or nervous
system. These phenomena may therefore be classed
with the reactive processes in the Annelida.
Thus a genealogical tree of inflammation can be
drawn up, starting with the researches on the reactive
phenomena of the invertebrata, and completed by facts
observed in the embryonic and early larval stages of the
vertebrata. These facts prove that the reactive pheno-
mena ensuing on lesions are in their origin essentially
the same in the two great branches of the animal
kingdom. But, whereas in the invertebrata the pro-
cesses have remained stationary, in the vertebrata they
have become in the course of development much more
complex in character. Even in the older larvae of
Triton and Axolotl, which are provided with a larger
number of blood-vessels wide enough to allow the
passage of leucocytes, the inflammatory reaction takes
place in the classical manner that has been so frequently
studied during the last twenty-four years. The same
lesions still produce first an acceleration, then a slowing
of the blood-stream, followed by an accumulation of
white corpuscles in the peripheral zone and their emi-
gration and movement towards the injured spot.
The tadpoles of the different Batrachians are well
adapted for the investigation of these phenomena ;
some, such as those of Bombinator igneus, are distin-
104 INFLAMMATION.
guished by their large fins, which are so transparent as
to admit of direct microscopic examination. If inflam-
mation be excited in the living tadpole by the applica-
tion of small fragments of silver nitrate, by a simple
puncture or by any other means, all the stages of in-
flammation and regeneration can be followed under the
microscope for several days or weeks in succession.
Hence the great advantage of these researches over
those prosecuted on the mesentery or tongue of the frog.
If we wish to fix any stage, we have only to adopt
Ranvier's method, 1 and drop the whole animal into 25
per cent, alcohol. After it has been for a few hours in
this liquid, the epidermis may be detached by means
of forceps, the tail cut off and covered with a few drops
of an aqueous solution ofvesuvine. After some minutes
the preparation is sufficiently stained, and may be
washed with water and examined just as it is, without
being dehydrated and mounted in balsam.
The tail of tadpoles shows marked anatomical differ-
ences compared with the fin of young larvae of Triton
or Axolotl. In the former the blood-vessels are much
more numerous and more richly branched, while the
amoeboid cells normally present in the connective
tissue are much less numerous. All the histological
elements, the cells of the connective tissue as well as the
red and white blood-corpuscles, are smaller. These
peculiarities in the tadpole favour the emigration of
leucocytes, which is very pronounced after the infliction
of the various lesions that I have practised, such as
section of the end of the tail, puncture by a foreign
body or cauterisation with nitrate of silver. Sometimes,
as soon as a quarter of an hour after the infliction of the
1 " TraW technique d'histologie," 2* edition, 1889.
LECTURE VII. 105
injury, we may see the beginning of inflammation,
accompanied by vascular dilatation and a considerable
emigration of leucocytes. In cases where the irritation
is more lasting, as after the introduction of a splinter,
the inflammatory emigration may be observed for
several days in succession. The result is an accumula-
tion of leucocytes round the seat of lesion, an accumula-
tion infinitely more extensive than that which occurs in
the young larvae of Triton and Axolotl. A comparison
of the reactive phenomena in these Urodela with those
in tadpoles of Batrachians demonstrates in the clearest
manner possible how much more marked the reaction
becomes when the vascular system co-operates.
It is very probable that the migratory cells of the
connective tissue also move towards the injured spot.
Their number, however, is so limited that they are quite
unimportant in comparison with the mass of emigrated
leucocytes. The fixed connective tissue cells undergo
the same changes that have been described in the case
of the Tritons. Immediately after the application of
the irritant, they swell up, become vacuolated and less
highly refracting, while their processes become shorter
and lose many of their branches. A short time later,
however, these cells resume their normal condition, in-
cluding their stationary position and other characteris-
tics.
The migratory cells which have collected round the
lesion in the young Urodela, as well as the numerous
leucocytes which have emigrated from the blood-
vessels in the tadpoles and in the older larvae of
the same Urodela, at once show marked phagocytic
properties. They greedily devour coloured granules
which have been either introduced on a foreign body
106 INFLAMMATION.
or simply rubbed into the wound ; they also englobe
granules of dead pigment-corpuscles or other cellular
debris.
In my first paper on inflammation in amphibia 1 I
laid stress on the fact that the fixed ramified cells of the
connective tissue are also possessed of phagocytic pro-
perties. On examining fins which have been inflamed
for several days we may readily observe cells containing
foreign bodies, such as granules of carmine or broken-
down red corpuscles, in their protoplasm, although these
cells have antler-like ramifications and must there-
fore be regarded as characteristic connective tissue
cells. From these facts, which I have confirmed several
times, I concluded that these cells were phagocytes,
just as much as the migratory cells. But I have since
recognised that this deduction was erroneous. In spite
of numerous attempts, I have never succeeded in
observing the inclusion of foreign bodies by the proto-
plasmic processes of the connective tissue cells. My
researches on this point, which I have repeated for
several years, have convinced me that the fixed cells,
once definitely formed, never under any circumstances
englobe granules of carmine or other substances. The
granules found enclosed in them have been taken up in
a previous stage of their development, when they were
still mobile phagocytes. These facts afford therefore
certain proof of the conversion of migratory cells into
fixed connective tissue cells. Although this conclusion
is opposed to the almost unanimous opinion of patho-
logists, it is nevertheless correct.
It does not, however, necessarily follow that all the
mobile cells which have collected at the seat of lesion.
1 Biologisches Centralblatt, 1883.
LECTURE VII.
107
are transformed into fixed branched cells. A great
number of phagocytes do not undergo this change ;
many of them perish and are englobed by other phago-
cytes, as can be seen in every case a few days after the
onset of the inflammation (Fig. 52). Several of these
wandering cells penetrate into the epidermis, and escape
thence to the surface, where they meet a certain death
in the surrounding water. Others again pass into the
lymphatics (PI. I. Fig. i) and are
carried away by the lymph-stream.
A certain number of phagocytes
remain at the seat of the lesion, in
spite of the fact that complete
regeneration may have occurred.
In the cases where the irritant
remains in the tissue, as for in-
stance the small glass tube which
has been introduced into the con-
nective tissue of tadpoles, a num-
ber of the emigrated leucocytes
remain at the injured spot, some-
times surrounding the foreign body
for several months.
This description of inflammation in the tadpole may
be taken as a type of the same process in the other
Vertebrates, including mammals and man himself.
We have thus arrived at the final complication of
the inflammatory reaction as it occurs in the animal
kingdom. Before examining this pathological process
in detail, it would be advantageous to cast a look
back on the evolution of this important phenomenon.
Since the chief cause of inflammation infection
Fig. 52. Phagocytes enclosing;
other phagocytes. From the
fin of a tadpole of Boinbi-
nator.
IO8 INFLAMMATION.
must be considered as a struggle between two organisms,
the parasite and its host, and since this struggle brings
about adaptations on both sides, we must admit that the
organism has elaborated means to defend itself against
its aggressors. If the unicellular beings, in which the
nature of the struggle is more evident, already possess
means of defence, we certainly cannot suppose that the
higher animals are deprived of equal powers.
The plants, which are incapable of motion, defend
themselves by the secretion of thick tough membranes,
to penetrate which the parasite has to adopt special
measures. Thus it must either secrete ferments which
dissolve the cellulose, or pierce the membrane by the
force of its growth, so that the plant is well defended
against a large number of parasites which are not
possessed of these powers.
The penetration of parasites into, or the action of any
other injurious agents upon vegetable cells causes their
death. But even if these cells are incapable of regenera-
tion, others, which remain intact or are even excited
by the irritant, multiply and take their place. In the
plant world, then, we have lesions and primary ne-
crosis ; regeneration also takes place, often to full re-
integration, but there is no such thing as inflammation.
The animal condition of some lower plants, especially
the plasmodium of Myxomycetes, offers an exception
to this rule, since here we have a vegetable organism
capable of locomotion and cellular digestion. The
latter property, which consists in taking up and digest-
ing or excreting the deleterious matter, contributes to
the protection of the organism. Although it may
not constitute inflammation itself, at any rate it repre-
sents a primitive condition of this process.
LECTURE VII. 109
Inflammation makes its appearance only in the
animal kingdom and undergoes a slow evolution, which
begins in the organisms that have a mesoderm. At
first it cannot be distinguished from a simple intra-
cellular digestion, effected by amoeboid and phagocytic
cells of the mesoderm. Thus in the Sponges the diges-
tive and the inflammatory functions are still united ;
but as soon as the endoderm becomes definitely
separated from the mesoderm, the two functions di-
verge. The endoderm now acts exclusively as a diges-
tive organ, while the mesoderm alone retains the power
of protecting the organism against injurious agents by
digesting them when possible. The mesodermic phago-
cytes preserve their property of intracellular digestion ;
this they effect either by fusing into plasmodia or by
collecting to form capsules round the parasites or other
foreign bodies. The phagocytic reaction is displayed
by all the mesodermic phagocytes. In this process
the prominent part is played in some cases by the
connective tissue cells, in others by the peritoneal cells
or by the corpuscles of the perivisceral fluid or of the
blood. In all these cases it is the phagocytes which
war against the aggressor by devouring, englobing and
digesting it.
It is apparent that the inflammation of vertebrates,
in which the defending phagocytes emigrate from the
vessels to proceed against offending bodies, is distin-
guished only quantitatively from the analogous phe-
nomena in invertebrates and must therefore be also
regarded as a reaction of the organism against dele-
terious agents. We must conclude that the essential
originating factor, the primuni movens of inflammation
consists in a phagocytic reaction on the part of the
HO INFLAMMATION.
animal organism. All the other phenomena are merely
accessory to this process, and may be regarded as means
to facilitate the access of phagocytes to the injured part.
The morbid phenomena properly speaking, such as
the primary lesion or necrosis, as well as the processes
of repair, do not form part of the inflammation and
must not be confounded with it.
The discussion as to which stage in the evolution of
the phagocytic reaction should have the name of in-
flammation applied to it, is really beside the point.
We might follow Strieker, 1 Roser, 2 and others, in limiting
this term to the phenomena in which the vessels co-
operate. In this case the phagocytic reaction of the
elder larva? of Urodela would fall under the head of
inflammation, while the same reaction in younger larva?,
where the phagocytes are furnished by the connective
tissue, would have no right to this term. If on the
other hand we wish to adhere to the etymological
meaning of the word, we can only apply the name of
' inflammation ' to those cases where the phagocytic
reaction is accompanied by increased local temperature,
that is to say, we must reserve it exclusively for the
warm-blooded animals. The important points, with
which we are concerned, are the recognition of the
natural relations between the phenomena and the
tracing of their genealogical history.
This general conclusion, to which we have been led
by a comparative survey of the phagocytic reaction,
will, I think, facilitate a closer study of inflammation
in the vertebrata.
1 " Allgemeine Pathologic der Infectionskrankheiten." Wien,
1886, p. 112.
- " Entziindung und Heilung.' 1 Leipzig, 1886, p. 55.
LECTURE VIII.
Varieties of leucocytes Origin of these varieties Mobility
Phagocytic properties Condition of englobed microbes
Their vitality and virulence Sensibility of leucocytes-
Tactile sensibility Chemiotaxis Buchner's investigations
Leucocytosis Intracellular digestion Destruction of mi-
crobes, especially in immune animals Action of leucocytes on
spores Multiplication of leucocytes by direct and indirect
cell-division Changes they undergo Transformation of lobed
into single nuclei.
FOR the purpose of simplification, I propose to treat
the various parts of the inflammatory reaction in verte-
brates separately, beginning with a study of the principal
actors in this process the leucocytes.
Of these cells, which include the colourless corpus-
cles contained in the blood and lymph, several varieties
may be distinguished. In the first place, we find small
leucocytes with a large round nucleus and a very
small amount of protoplasm forming a thin layer round
the nucleus. These cells (Fig. 53^) which are often
spoken of as lymphocytes because they occur in large
quantities in the lymphatic glands, stain readily with a
variety of colours, especially with the aniline dyes. The
latter stain the nucleus deeply, and the protoplasm only
faintly. The lymphocytes gradually grow until they
become large-sized leucocytes, provided with a single
nucleus, which contains a considerable proportion of
112 INFLAMMATION.
nucleoplasm and is easily stained with the aniline
colours. The nucleus of these large leucocytes, how-
ever, does not stain so deeply, whereas their protoplasm
stains as well as or better than that of the lymphocytes.
Among these mononuclear leucocytes are some cells with
a round or oval nucleus (Fig. 53^) and others with a
kidney- or bean-shaped nucleus. This kind of leucocyte
bears a great resemblance to certain fixed connective
tissue cells as well as to endothelial cells and cells of
the splenic pulp. On this account it is often difficult to
distinguish the mononuclear leucocytes from these cells,
especially when they
occur outside the
vessels.
The two next
varieties of leuco-
cytes on the other
hand may easily
be recognised, even
Fig. 53- Three forms of leucocytes.
when surrounded
with all sorts of histological elements. The first of
these, the eosinophile leucocytes of Ehrlich, are provided
with a nucleus, generally lobed and of very variable
form. They contain in their protoplasm coarse granules
which do not stain with the basic aniline dyes, such as
fuchsine, methyl or gentian violet, methylene blue,
vesuvine and others, but readily take up the acid
aniline colours, especially eosin, which stains them a
rose-red. (Plate III. Fig. 2.)
The fourth variety of leucocytes, the most important
quantitatively and even qualitatively, is represented by
cells in which the nucleus is either lobed or composed
of several portions united by such delicate nuclear fila-
LECTURE VIII. 113
ments (Fig, 53*:) as to give the impression of a multi-
nucleated cell. Most of the nuclei are in this form,
although they are sometimes really multiple, thus
justifying the name of polynnclear leucocytes, which has
been given to this kind. The shapes of these com-
pound nuclei are very variable. They more frequently
resemble a trefoil or a raspberry, but they may occur in
the form of a ring. Besides the nucleus these leuco-
cytes possess an attraction-sphere composed of fine fila-
ments of achromatin and containing a small central
body of chromatin. This peculiarity has been lately
discovered by Flemming * in the leucocytes of salaman-
der larvae. (Plate II. Fig. 7.)
The polynuclear leucocytes differ from the other
colourless corpuscles in their reaction to aniline dyes,
which stain the nuclei very deeply, while the protoplasm
remains for the most part unaffected. The latter con-
tains granules, sometimes, as in man, in great number.
These can only be stained by a mixture of the acid with
the basic dyes, so that these polynuclear leucocytes are
often alluded to as neutrophile leucocytes. 2
Although the fact that there are several kinds of
leucocytes was shown by Max Schultze 3 as long ago as
1865, an exact knowledge of the different forms dates
only from the discoveries of Ehrlich. 4 On examining the
relative proportion of the various leucocytes in the blood,
1 Archiv fur mikroskop. Anatomic, 1891, vol. xxxvii. p. 249.
Plates 1 3 and 14.
2 In the rabbit and guinea-pig these cells are replaced by
amphophile or pseudoeosinophilc leucocytes.
3 Archivf. mikroskop. Anatomic, 1865, vol. i.
4 The valuable papers of Ehrlich have been recently collected in
a single volume. " Farbenanalytische Untersuchungen zur His-
tologie und Klinik des Blutes." Berlin, 1891.
I
1 14 INFLAMMATION.
it is found that the neutrophile cells are by far the most
numerous, forming three-fourths of the total amount.
It was at first thought that these varieties depended
on a difference in the origin of the leucocytes and that
the small cells were formed only in the lymphatic
glands, while the others were derived from the bone-
marrow. Leucocytes were accordingly divided into
two groups, lymphocytes and myelocytes. This classifica-
tion had to be abandoned for two reasons. In the first
place, small cells can also be formed in other organs,
such as the spleen and the bone-marrow ; and, in the
second place, the myelocytes do not originate solely in
the bone-marrow. Here again our most accurate know-
ledge is due to Ehrlich, who has shown that the eosino-
phile cells are produced specially by the bone-marrow;
hence an abnormal number of these cells in the blood
serves to indicate that this tissue is primarily affected,
as in myelogenic leuchaemia. The ordinary polynuclear
leucocytes or neutrophile cells develop in the blood itself
at the expense of the small cells which have been formed
in various organs. This fact has been specially pointed
out by Ouskoff. 1
In considering this question of the origin of leuco-
cytes, it must not be forgotten that these cells are to
be found in the blood of the inferior fishes, such as
Cyclostomata, which possess neither lymphatic glands,
bone-marrow nor spleen, and in which the leucocytes
are derived from the mesodermic cells of the embryo
and perhaps also from the endothelium of the blood-
vessels.
The leucocytes, as Lieberkiihn showed, are mobile
cells which, like the amoebae, have the power of putting
1 "Le sang comme tissu," (Russian), St. Petersburg, 1890.
LECTURE VIII. 115
out protoplasmic processes and of moving from place
to place. All the leucocytes have this property ; but
it is not so well developed in the lymphocytes, which
are the youngest of the white blood-corpuscles. Neither
these nor the eosinophile cells are able to englobe
foreign bodies, and therefore cannot act as phagocytes^
Hence it is probable that the granules, which are so
characteristic of the eosinophile cells and in reptiles
and birds are shaped like small rods or crystals, are not
taken up by the cells, but are manufactured in the body
of the cell itself. On the other hand the two remain-
ing varieties of leucocytes, the mononuclear and the
neutrophile corpuscles are endowed with very pro-
nounced phagocytic properties. Even outside the
organism these amoeboid cells readily englobe a large
number of foreign particles with which they may come
in contact, and they maybe often seen literally crammed
with all sorts of granules. Like the amoebae, they
swallow not only inert bodies, such as granules of
carmine or other substances that are insoluble in the
fluid surrounding the leucocytes, but also a large num-
ber of living organisms. Thus the leucocytes of the
frog englobe the bacilli which cause a septicaemia in
these Batrachians. That the bacilli are in a living
condition when swallowed, is shown by the fact that
they perform active movements although enclosed in
the nutritive vacuoles of the leucocytes. 1 The same
thing is observed if we introduce leucocytes filled with
bacteria into a medium which nourishes the bacilli
while killing the leucocyte. Thus on introducing the
leucocytes of the pigeon filled with anthrax bacilli (to
which the pigeon is very refractory) into bouillon, the
1 Vide Biologisches Centralblatt, 1883, p. 562.
1 2
Il6 INFLAMMATION.
bacilli grow, pierce the protoplasm of the cells, and
form well-developed filaments, 1 showing definitely that
the bacilli were englobed in a living condition. This
has also been proved in the case of other bacteria.
Thus the Vibrio Metchnikowii that has been taken up
by the leucocytes of immune guinea-pigs, will develop,
even in an exsudation obtained from the animal itself, if
the fluid be put under conditions unfavourable to the
life of the animal cells. 2 The conclusion is further con-
firmed by the fact that masses of bacilli of tuberculosis,
of swine septicaemia (' rouget des pores ') and of mouse
septicaemia are'englobed by the leucocytes of a large
variety of animals whether susceptible or refractory to
these diseases.
But although it may be clearly shown that the leuco-
cytes enclose living microbes, we must not conclude
that these cells devour all microbes indiscriminately.
Thus in many cases we find that the leucocytes of an
animal which is very susceptible to a certain kind of
bacteria, do not take up these bacteria at all, even
though they may be in contact with them. As an
example we may quote the behaviour of the leucocytes
of mice and guinea-pigs towards the bacilli of anthrax,
or that of the leucocytes of pigeons and rabbits towards
the bacteria of chicken cholera, or of the leucocytes of
guinea-pigs that are susceptible to vibrionic septicaemia
when brought in contact with the exciting agent of this
disease.
It is evident then that leucocytes can englobe viru-
lent microbes.' This fact may be shown in another
way, The two kinds of leucocytes which 'are phago-
1 Annales de I'lnstitut Pasteur, \ 890, p. 80.
* Ibid., 1 89 1, p. 471.
LECTURE VIII. II/
cytic in their action, the large mononuclear and the
neutrophile leucocytes, vary in their behaviour towards
different species of microbes. Thus in man the mono-
nuclear leucocytes do not take up either the strepto-
coccus of erysipelas or the gonococci, whereas these
two microorganisms are readily englobed by the neu-
trophile polynuclear leucocytes. 1 This choice shows
that the microbes thus avoided by the mononuclear
leucocytes, are by no means inactive bodies, otherwise
they would be taken up by this variety of leucocytes,
just like any other inert matter. Leprosy bacilli, on the
other hand, are never englobed by the neutrophile poly-
nuclear leucocytes, but are readily devoured by the
mononuclear cells. 2
These differences must be due to a varying sensibility
to microbes on the part of the two kinds of leucocytes,
dependent no doubt in both instances on chemio-
taxis.
Some researches on phagocytosis in amreboid cells 3
suggested that action at a distance must play some
part in these phenomena. We owe to Leber, 4 however,
the first clear exposition of the part taken in them by
the chemiotactic sensibility of the leucocytes. In his
experiments on keratitis produced by a crystalline
1 Virchow's Archiv, vol. cvii., 1887, p. 227.''
2 Ibid., p. 228, and Savvtchenko, Ziegler's Beitr'dge znr patholo-
gischen Anatomic, vol. ix., 1890, p. 252.
3 " Untersuchungen iiber die intracellulare Verdauung," in the
Arbeiten des zool. Inst.zu Wien, 1883, vol. v. p. 159.
4 Fortschritte der Median, vol. vi., 1888, p. 460.' See also Leber's
large work, " Die Entstehung der Entziindung," published at
Leipzig in May, 1891. As my manuscript was already written when
this monograph appeared, I have not been able to quote from it so
often as I should have liked.
IlS INFLAMMATION.
substance extracted from cultures of Staphylococcus
aureus, he showed that the leucocytes at a distance
were attracted towards the point where this substance
had been introduced. On putting some small glass
tubes filled with this substance, into the anterior cham-
ber of the eye, they became filled with a mass of leuco-
cytes, although the tubes were so placed that the cells
had to move against gravity in order to effect an entry
into them.
This important discovery was the starting-point of a
series of researches which proved beyond question the
existence in leucocytes of chemiotactic properties
absolutely analogous to those of plasmodia and other
lower organisms. Lubarsch 1 showed that the leuco-
cytes of the frog are more readily attracted by living
bacilli than by the same bacilli if previously killed by
heating. Pekelharing " then pointed out that the leuco-
cytes of this animal are much more strongly attracted
by the bacilli of anthrax than by a passive substance
such as cotton fibres. In this connection many facts
of great importance have been brought forward by
Massart and Bordet, 3 who proved that the leucocytes of
the frog are attracted by a number of culture fluids of
various microbes, especially of the Staphylococcus
fyogenes albus, as well as by the pleuroperitoneal
transsudation of frogs that have been poisoned by ox-
bile.
Among the products of the oxidation of albumen,
1 Fortschritte d. Medicin, 1888, vol. vi. No. 4, and Centralblatt
fur Bacteria logic, vol. vi. Nos. 18-20.
3 Semaine niedicalc, No. 22, 1889, p. 184.
3 " Recherches sur 1'irritabilite des leucocytes." Journ. publ.
par la Soc. dcs Set. med. et nat. dc Bnixelles^ Feb. 3rd, 1890.
LECTURE VIII. 119
examined by these authors, leucin was the only one
found to exercise an attractive influence on the frog's
leucocytes, while others such as creatin, creatinin,
allantoin &c., did not give rise to any chemiotaxis.
Gabritchevsky, 1 in a research carried out in my
laboratory, has shown that the leucocytes of mammals,
especially of the rabbit, are much more sensitive to
chemical excitation than are those of the frog. He
pointed out moreover that whereas leucocytes are
strongly attracted by sterilised or living cultures of
most pathogenic and saprophytic bacteria as well as
by papayotin, they are repelled by the most virulent
bacteria, such as those of chicken cholera, and by lactic
acid, ten per cent, solutions of sodium and potassium
salts, alcohol, chloroform, glycerine, jequirity, bile and
quinine. The leucocytes remain unaffected by many
other substances such as water, weak solutions of
sodium or potassium salts, peptone, phloridzin, &c.
Buchner,- after confirming these facts on the chemio-
tactic properties of leucocytes, has endeavoured to de-
termine more precisely the nature of the substances
contained in bacterial cultivations which call these
properties into play. In conjunction with Lange and
Romer he found that the proteins of various species
(bacillus pyocyaneus, bacillus subtilis, the typhoid bacillus,
Staphylococcus pyogenes aureus and others) exercised an
attractive influence on the leucocytes of the rabbit. He
obtained the same effect with solutions of gluten-casein,
as well as with some alkali-albumens of animal origin.
Buchner considers that these experiments warrant the
conclusion that it is only the contents of bacteria, and
1 Annales de flnstitut Pasteur, 1890, p. 346.
* Berliner klinische Wochenschrift^ 1890, No. 47.
120 INFLAMMATION.
not the products of their secretion, which exert a
chemiotactic influence on the leucocytes. We must
observe, however, that Buchner has not by any means
proved his contention. It is evident that the toxic
products of the bacteria must play the most important
part in evoking the phenomena of chemiotaxis. Now
it is precisely these products which adhere most strongly
to albuminoid and other bodies, so that we are still far
from being able to isolate them and study their effects
in a pure condition. On the other hand, the more
advanced products of disintegration, such as ammonia
and its salts, are not of such great importance in the
question of leucocytic sensibility in the animal body as
to permit of our drawing conclusions from the fact that
they exert no chemiotactic influence. And yet, even
among these substances, we find a body, leucin, which
has the property of attracting leucocytes. This fact
was observed by Massart and Bordet, and has been
confirmed by Buchner himself.
Physiological chemistry is not yet sufficiently ad-
vanced to give a decision in the question raised by
Buchner. We have not sufficient evidence therefore to
justify us in accepting the dictum laid down by this
author, viz. that the chemiotaxis of leucocytes can only
be excited by dead or injured bacteria, since only under
such conditions are their contents dissolved out into
the surrounding fluid. Even if Buchner's hypothesis
could be proved, we must remember that in every
medium tenanted by bacteria, there are a certain number
of dead ones among the living. The former would at-
tract the leucocytes, which would thus be brought in
contact with the living bacteria, and the final result
would be the same as if the living bacteria had them-
LECTURE VIII. 121
selves exercised the chemiotactic influence. But besides
these considerations, there are other facts which tell
against this hypothesis, as, for instance, the greater
attractive power possessed by living bacteria on leuco-
cytes, as shown in the experiments of Lubarsch.
Further there are the still more important cases in which
the bacteria or other parasites have been englobed by
the leucocytes in the living condition. Moreover the
evidence brought forward by Buchner himself argues
against his idea that the leucocytes are not attracted by
active bacteria. In experiments made in conjunction
with Romer, Buchner found that injections of proteins
of bacteria, especially of the bacillus pyocyaneus, into a
vein, produced a pronounced general leucocytosis, the
number of white corpuscles in the blood being seven
times the normal amount. Now leucocytosis is a con-
dition of extremely frequent occurrence in the infectious
diseases generally. Although in some, such as typhoid
fever in man, the number of leucocytes has not always
been observed to be increased, yet in the vast majority
of the other infectious disorders this has been found to
be the case. Thus several observers * have shown that
a pronounced leucocytosis accompanies infection by
anthrax in animals which die of this disease (such as
guinea-pigs, horses, oxen and others), although masses
of actively living bacteria occur at the same time in the
blood.
According to von Limbeck ~ and Pee, 3 leucocytosis
1 Bellinger, " Milzbrand," 1872, pp. 2, 101.
2 Von Limbeck, " Klinisches und Experimentelles iiber die ent-
ziindliche Leucocytose," Prag, 1889.
3 H. Pee, " Untersuchungen iiber Leucocytose," Berlin, 1890,
P 13-
122 INFLAMMATION.
is a constant phenomenon in erysipelas in man. It is
at its height during the febrile period, while the blood
contains a number of living streptococci, and comes
to an end after the crisis, when only masses of dead
microbes are to be found in the organism.
This coincidence in time of the leucocytosis and the
most active condition of the bacteria has been also
shown by von Limbeck in the case of fibrinous pneu-
monia in man. The number of the colourless cor-
puscles present at any time in the blood corresponds
exactly with the temperature ; when the disease ter-
minates in crisis, the leucocytes decrease suddenly,
while in the cases where the temperature drops slowly,
(lysis), the fall in the number of leucocytes is also
gradual.
Working with dogs, von Limbeck pointed out that
the injection of Staphylococcus pyogenes aureus into the
knee-joint was almost immediately followed by a large
increase of the leucocytes in the blood, before any
local symptoms had made their appearance.
We thus see that the special instance of chemiotaxis
known as leucocytosis, occurs at a time when the
greatest proliferation of microbes is taking place and
that the leucocytosis begins to diminish directly the
pathogenic bacteria die. We must remember also that,
according to other researches of Buchner, J these same
proteins, which he considers to exist exclusively in the
interior of the bacterial cell, are the active agents in
producing fever, that is, the symptom which occurs
while the pathogenic microbes are actively multiplying,
and ceases with their death.
Like the plasmodium of Myxomycetes and many other
1 Berliner klinische Wochenschrift, 1890, No. 30, p. 673.
LECTURE VIII. 123
inferior organisms, the leucocytes have other forms of
sensibility besides that of chemiotaxis. Their tactile
sensibility, which serves them in the inclusion of foreign
bodies, is very highly developed. When the leucocytes
meet a resistant surface, they react by offering as large
a surface of contact as possible (Massart and Bordet).
By means of this property, the leucocytes are able to get
through the finest pores, and can penetrate elder-pith
and even such compact bodies as bone and ivory. The
physical changes in the surrounding medium, such as
variations in temperature, pressure, or rapidity of move-
ment of the liquid, and probably many others, must be
appreciated by the leucocytes. It would be very in-
teresting to make a systematic study of these physio-
tactic properties of the white corpuscles.
Leucocytes which, by virtue of their chemiotaxis,
are attracted at a distance by microbes and other
particles, move towards these bodies and on coming in
contact with them englobe them owing to their tactile
sensibility. The ingested particles are now acted on
by the leucocyte. It is an old observation that red
corpuscles, when enclosed by leucocytes, partially dis-
solve, leaving a pigmented residue. It is also very easy
to follow the changes undergone by pus-corpuscles in
the interior of leucocytes, where they gradually lose
their staining power and are finally converted into
scattered granules which are partially dissolved.
These changes are carried out by the protoplasm of the
leucocytes, and must be looked upon as an act of intra-
cellular digestion. This view is justified by the dis-
covery of ferments in leucocytes. Thus Rossbach ' has
1 Deutsche meuccin. IVvchcnschrift, 1890, p. 389.
124 INFLAMMATION.
shown the existence of an amylolytic ferment in leuco-
cytes from various organs, especially from the tonsils,
and Leber l has proved that the pus from a hypopion,
entirely freed from microbes, will digest coagulated
fibrin at a temperature of 25. The same pus has also
the power of liquefying gelatin. This digestive power
can be destroyed by heat, showing that in this respect
the ferment of the leucocytes resembles all other fer-
ments.
The digestion of proteid substances by the leuco-
cytes is well shown by the gradual changes that take
place in the muscular fibres which have been englobed
by leucocytes in cases of acute muscular atrophy. The
presence of peptone in leucocytes, which has been so
often proved by Hofmeister, is sufficiently accounted
for by this fact of intracellular digestion and need not
therefore be referred, as done by this author, to an ab-
sorption by these cells of the peptone formed in the
alimentary canal.
Our knowledge of the conditions under which this
intracellular digestion takes place is still very incom-
plete. In a large number of experiments that I have
made on the absorption of granules of blue litmus by
leucocytes, I have seen the colour change to red only in
a few exceptional cases. ~ It appears then that diges-
1 " Die Entstehung der Entziindung,'' p. 508.
3 Annales de PInstitut Pasteur, 1889, p. 29. Netchaeff ( Virchoufs
Archiv, vol. cxxv., 1891, p. 448) thinks that in the cases where I
observed a conversion of the blue into red granules, the change was
simply an optical illusion. This he concludes from his own ob-
servations on the fate of litmus granules in the interior of leucocytes.
My researches, which extend over a series of years, leave me,
however, in no doubt as to the reality-of the change in colour of the
litmus. I must add that the criticism by Netchaeff of my work
LECTURE VIII. 125
tion is carried out by leucocytes in a neutral or alkaline
medium, as in the case of the phagocytes of the
Sponges.
Not only are the disintegrated muscle fibres or blood
corpuscles digested by the leucocytes, but in a large
number of cases the microbes that have been devoured
also undergo the same fate. This digestion of bacteria
in the interior of leucocytes can be most readily studied
in refractory animals. Nothing is more instructive to
watch than the changes undergone by the streptococci
of erysipelas in the leucocytes of white rats. The bac-
teria that have been englobed fuse into irregular masses
which do not stain completely, a process closely re-
sembling the digestion of the sulphobacteria in the
body of Stentor, discussed in the second lecture.
Anthrax bacilli are digested in the same way by the
leucocytes of a number of immune animals. The frog's
white corpuscles, instead of presenting a favourable
medium for the development of anthrax bacilli as
Koch l and Petruschky - have asserted, not only prevent
their growth and development, but also kill and digest
these bacteria. All the phases of this digestion corre-
spond exactly with those of the digestion of the sapro-
phytic bacilli by amoebae (vide Chap. II,). As in this
case, a number of the bacilli in the interior of the
leucocytes are stained deeply by an old aqueous solu-
tion of vesuvine, which also colours other particles that
are being digested by the amoebae, as Bruno Hofer has
on intracellular digestion shows that this observer has never
examined the most classical examples of this process, more espe-
cially the digestion in Protozoa.
1 " Beitriige zur Biologic der Pflanzen," published by Cohn, 1876,
vol. ii. p. 300.
Zcit'ichrifl fiir Hygiene, 1889, vol. vii. p. 75.
126
INFLAMMATION.
also shown. This period during which the devoured
bacilli are deeply stained, is followed by a stage in
which the bacilli are only partially stained, and finally
even this property is lost and they remain absolutely
uncoloured. At last only the more durable cell-mem-
brane of the microbe is to be seen in the leucocyte, and
even this disappears at the end of a cer-
tain space of time. The bacilli which
have been englobed by leucocytes are
much more rapidly digested in the case
of mammals that are either naturally
refractory, as the dog and fowl, or
have been rendered artificially immune
against anthrax by vaccination, as the
rabbit. This fact is shown by the
researches of Hess 1 as well as my
own. 2
It is easy to follow the digestion of
many other microbes within the
leucocytes. Vacuoles are often seen
to form around the bacteria that have
Fig. 54.-An anthrax been swallowed, just as we have
bacillus, stained by ...
vesuvine, in a leucocyte noticed in the digestion of nutrient
of the frog. The two
figures represent two material by the protoplasm of the
phases of movement of
the same ceil. Protozoa and the Myxomycetes. I
have been able to observe the changes undergone by
the spirilla of recurrent fever in the leucocytes of
monkeys/' as well as those undergone by the vibrio of
septicaemia in the leucocytes of immunised guinea-pigs,
and those by erysipelas streptococci in the leucocytes
1 Virchoitfs Archiv, 1887, vol. cix. p. 365.
2 Ibid., 1884, vol. xcvii. p. 502.
* Ibid., 1887, vol. cix. p. 176.
LECTURE VIII.
127
of man, 1 &c. We are at present ignorant of the precise
manner in which this digestive and destructive action
is accomplished, and do not even know whether the
substance which kills the microbes is a ferment or not.
The fact that the ferments of the higher animals, such
as pepsin and trypsin, do not kill bacteria, is no reason
frig- 55 Four leucocytes from the frog, enclosing anthrax bacilli. Some are alive
and unstained, others which have been killed have taken up the dye.
for assuming that there may not be other ferments
which are capable of exercising a bactericidal action.
Some authors have denied to leucocytes the posses-
sion of digestive powers. Schafer 2 comes to this con-
clusion from the observation that neither proteid sub-
1 Virchovfs Archiv, vol. cvii. p. 209.
* British Medical Journal, 1882, No. 1134, p. 573.
128 INFLAMMATION.
stances nor granules of fat or starch underwent any
change when ingested by the leucocytes of Tritons.
But as his researches were carried out on leucocytes
that had been removed from the body and immersed in
normal salt solution, they cannot be taken to refute the
facts that have been acquired from a study of the living
animal. Netchaeff has recently endeavoured to disprove
the intracellular digestion of bacteria by leucocytes, but
we have already shown how inadequate are the argu-
ments he brings forward.
It is undeniable then that leucocytes possess digestive
powers, and that in particular they are able to digest
microbes. But it does not therefore necessarily follow
that these cells kill and digest all the microbes they
englobe. In certain diseases the leucocytes take in a
number of bacteria, such as tubercle bacilli or the
bacilli of" swine erysipelas or mouse septicaemia, a few
of which may be digested while the others resist the
digestive action of the leucocytes, multiply in the cells
and finally invade the whole organism.
In several cases where the leucocytes have not the
power of killing the microbe, they can nevertheless
hinder it from growing and exerting its toxic influence.
The best examples of this arc afforded by bacterial
spores, which are endowed with great powers of resis-
tance. Anthrax spores are readily ingested by the
leucocytes of many species of animals, and, among
others, by those of immune animals such as the frog and
fowl. In spite of the fact that the spores germinate
and grow easily in the lymph-plasma of the two last-
mentioned, they are incapable of doing so in the bodies
of the leucocytes themselves, where they cannot exert
their injurious effects, although they preserve their
LECTURE VIII. 129
vitality and their virulence often for a considerable
length of time. Once, however, that the vitality of these
cells has been extensively lowered, as may be effected
in fowls by cooling them or in frogs by heating them
artificially, the spores germinate within the dead or en-
feebled leucocytes, and infect the whole body. 1 These
experiments were repeated several times by Trapez-
nikoff 2 in my laboratory, and show in a most striking
manner the important and salutary part played by the
leucocytes, in cases where these cells are unable to kill
the spore, and where the fluids of the organism are
totally inadequate to protect the infected animal.
The proliferation of leucocytes is at present the sub-
ject of much discussion. The polynuclear forms, with
fragmented nucleus, can give rise to two new leucocytes
by mere division of their protoplasm. This pheno-
menon has been described by Ranvier 3 and observed by
several other workers, especially by Arnold. 4 The fact
of the occurrence of simple or direct division of poly-
nuclear leucocytes has at length been generally accepted,
although for a long time after the discovery of karyo-
kinetic or indirect cell division, it was considered doubt-
ful, and all cells were supposed to divide in the latter
way. The phenomenon of direct cell division once
established, people have gone to the opposite extreme
and denied to the polynuclear leucocytes the power of
dividing in any other manner. All the cases in which
karyokinetic division of leucocytes had been observed,
1 See Wagner's paper on this subject, Annales de Flnstitut
Pasteur, 1890, p. 570.
2 Ibid., 1891, p. 362.
3 " Traite" d'histologie technique," 2nd edition, 1889, p. 137.
4 Archivfur mikroskopische Anatomic, 1888, p. 270.
K
130 INFLAMMATION.
from those of Peremeschko and Koultchitsky 'onwards,
were looked upon as open to doubt. Yet it has several
times been proved that leucocytes are capable of dividing
indirectly. Flemming 2 has observed this phenomenon
on several occasions, and has recently confirmed it
in the case of the leucocytes of the salamander.
Spronck, 3 too, has seen mitotic division of the leuco-
cytes in the blood of the rabbit and has found that at
any given time about 2. per 1000 of these cells are nor-
mally undergoing indirect cell-division. I can com-
pletely endorse this statement, since I have observed
undoubted mitotic division of the rabbit's leucocytes in
preparations made by Dr. Muskatbluth in my laboratory
at Odessa. I have on several occasions noticed the
same phenomenon in the migratory cells of the larvae of
Axolotl. In all the above cases it is the large mono-
nuclear leucocytes that divide indirectly, while the
polynuclear multiply by direct cell-division. All the
phases of the karyokinetic division of the leucocytes
that have emigrated from the vessels, and the conversion
of the new cells into amceboid cells may be studied in
the caudal fin of the living larvse of the white Axolotl.
(Fig. 56.)
In the same way it has been .denied that the leuco-
cytes possess the power of gradual transformation. The
fact that numbers of these cells are found dead in
exsudations has given rise to the idea that leucocytes
are in most cases destined to perish and are incapable of
forming constituent cells of any tissue. The lobed
shape of the nucleus in the polynuclear leucocytes has
1 Centralblatt fur medicinische Wissenschaften.
2 Archiv fur mikroskopische Anatomie, vol. xxxvii., 1891, p. 249.
3 Nederlandsch Tijdscrift voor Geneeskunde, March 29th, 1889.
LECTURE VIII.
even been taken as a sign of disintegration. Under
the influence of these views, many authors would refuse
to the leucocytes the power of conversion into fixed
connective tissue cells or even into epithelioid or giant
cells. In fact they now hold the exact opposite of the
theory formerly maintained by Cohnheim and especially
by Ziegler. 1 The latter observer at first regarded the
granulation-cells as well as the epithelioid and giant
cells of tubercle as derived exclusively from mononuclear
leucocytes. He has now abandoned this position and
Fig. 56. A migratory cell from the fin of a larva of Axolotl in process of division
by karyokmesis.
has adopted the opinion of his former opponents, who
believe the leucocyte to be incapable of giving rise
to any other form of cell. At the International Con-
gress held at Berlin in 1890, Ziegler, 2 relying chiefly on
researches that had been made by Nikiforoff in his
laboratory at Freiburg, stated that " the leucocytes do
not take any active part in the new formation of tissues."
1 " Ueber die pathologische Bindegewebsneubildung," 1875.
2 Centralblatt fur allgemeine Pathologic, 1890, Nos. 18 and 19,
P 575-
K 2
132 INFLAMMATION.
Marchand l and Grawitz - quoted their own original
observations in confirmation of this view. According
to them the leucocytes that emigrate during an inflam-
mation are not converted into granulation-cells, but
perish in the exsudation, and are either absorbed by the
lymphatics or devoured by amreboid cells derived from
the fixed connective tissue cells. Ribbert s holds a
similar opinion.
NikiforofFs 4 experiments, on which Ziegler chiefly
bases his view, were made on the subcutaneous con-
nective tissue of dogs, in which the formation of
granulation-tissue was provoked by the introduction
of drainage tubes. Although he affirms that this re-
generation of tissues is due exclusively to the pro-
liferation of the tissue-cells, and that the leucocytes
do not take any more active part in this process
than the red corpuscles or the fibrin, yet he himself
confesses he is unable to bring forward adequate proof
that the "cells originating from the blood-vessels are
incapable of further development and of conversion into
epithelioid cells and fibroblasts." 5 NikiforofT adds :
" Judging from my observations, I am inclined to the
view that in the granulation-tissue the mononuclear leu-
cocytes are converted by fragmentation of the nuclei
into polynuclear forms and are then taken up by
phagocytes." 6 This sentence is certainly much less
positive in tone than his conclusion quoted above and
than Ziegler's dictum. If we read Nikiforoff's account
of the observations themselves, we shall see that they by
1 Centralblatt fur allgemeine Pathologic, 1890, p. 577.
2 Ibid., p. 578. 3 Ibid., p. 665.
4 Ziegler's Beitmge zur pathologischen Anatomic, vol. viii., 1890,
p. 400.
5 Ibid., p. 415. c Ibid., p. 421.
LECTURE VIII.
133
no means prove his principal thesis, i.e. the non-par-
ticipation of the leucocytes in the formation of granula-
tion-tissue. The appearance of granulation-cells at a
period when the fixed connective tissue cells are in
process of multiplication, does not afford any proof that
the leucocytes are not also involved. Another argu-
ment brought forward by this author, the occurrence of
mitotic division, does not
in the least support his
conclusion, since we now
know that the leucocytes
are undoubtedly capable
of this mode of division.
It is not a matter for
surprise that Nikiforoffs
results should be want-
ing in accuracy and
clearness. The material
he worked with, the
granulations of the sub-
cutaneous tissue in dogs,
is not well adapted for _.
Fig. 57. Transition stages between amoeboid
the investigation of the jSJjJSj* From the tail of a tadpole of
processes under discus-
sion. In order to obtain reliable results, we must make
use of a subject on which we can follow the course of
events from day to day. For instance if we take the tail
of a batrachian tadpole and injure it in any way and watch
it for several days or even weeks in the living condition,
as I have done in my researches, it may readily be seen
that in these animals the polynuclear cells are converted
by a fusion of their nuclei first into mononuclear and
then into typical branched connective tissue cells.
134
INFLAMMATION.
(Fig. 57.) In order to bring out this fact more clearly
it is advisable to inflict the wound with an instrument
YXicJuLtl
Fig. 58. Inflamed spot of the tail of a larva of Bombinator igiieus 8 days after
cauterisation.
powdered with carmine. The granules of this substance
which have been ingested by the emigrated leucocytes,
are found in the interior of the connective tissue cells, the
LECTURE VIII. 135
nature of which is easily recognised by their charac-
teristic antler-like processes. (Fig. 58.)
The view accepted at the Berlin Congress cannot be
maintained. It is not the new theory of Ziegjer, but
his old way of thinking of 1875-1876 which is correct.
Although my personal observations have only been
concerned with amphibia, yet they agree perfectly with
the facts that have been ascertained in the higher verte-
brata by all observers, not excepting Nikiforoff himself.
In further support of my view I may quote the last
communication of Flemming, 1 who has also seen the
lobed or multiple nuclei of the migratory cells become
fused to a single rounded nucleus in the gills of sala-
mander larvae.
In the mammals, however, we have no proof that the
polynuclear can be transformed into mononuclear
leucocytes. Although the latter do certainly become
converted into epithelioid and giant cells, the poly-
nuclear variety seems unable to be thus transformed.
I need however only adduce the fact of the formation
of epithelioid and giant cells at the expense of mono-
nuclear leucocytes in the interior of the vessels in
rabbits which have received an intravenous injection
of tubercle bacilli. This phenomenon, which was de-
scribed by Yersin, 2 has been likewise observed several
times by myself. 3 Tchistowitch 4 in a research carried
out in my laboratory, has also witnessed all the
transition stages between typical mononuclear leuco-
1 Archiv f. mikroskop. Anatomic. Vol. xxxvii., 1891, p. 277.
2 Annales de F Institut Pasteur, 1888, p. 257.
3 Virchovfs Archiv, July, 1888, p. 88.
4 Annales de flnstitut Pasteur, July, 1889, p. 347, and PI. vi.
Figs. 5 and 7.
136 INFLAMMATION.
cytes and epithelioid and giant cells in the pulmonary
alveoli of rabbits.
In summarising this review of the present state of our
knowledge on the subject of leucocytes, we may con-
clude that the two varieties of these cells which play the
principal part in inflammation the mononuclear leuco-
cytes and the neutrophile cells are elements endowed
with very considerable chemiotactic and physiotactic
sensibilities ; that they are capable of amoeboid move-
ments, and prone to englobe and digest different foreign
bodies, especially a number of living microbes. In the
amphibia at least, the polynuclear can be converted into
mononuclear leucocytes and can become fixed connec-
tive tissue cells. In the vertebrata generally, the mono-
nuclear leucocytes may undergo transformation into
epithelioid and giant cells.
All that has been said on the subject of leucocytes
refers equally to all migratory cells.
LECTURE IX.
Endothelium of vessel walls Their development out of the mobile
cells of the embryo Development of the capillaries Contrac-
tility of the endothelial cells Star cells Phagocytosis in the
endothelial cells Fixed connective tissue cells Clasmatocytes
of Ranvier Ehrlich's cells Active migration of leucocytes in
diapedesis Experiments with quinine (Binz, Disselhorst)
" Itio in partes " Dilatation of the vessels Theory invoking
the influence of the surrounding tissue Influence of the
nervous system Negative chemiotaxis of the leucocytes in
cases of severe infection.
NEXT to the leucocytes, the vessels and their endothe-
lial lining play the most important part in inflam-
mation. According to modern embryological research
on fishes l the first endothelial cells of the vessels are
formed from the mobile cells on the surface of the
yolk sac. It is therefore not a matter for wonder that
the endothelial elements should have retained a few
traces of their ancient power of movement. The at-
tention of investigators has been especially directed
towards the protoplasmic processes of the endothelial
cells of the vessels during the new formation of capil-
laries. Conical buds of protoplasm make their ap-
pearance on the vascular walls. A protuberance show-
1 Vide H. Ziegler, " Die Entstehung des Blutes der Wirbelthiere. 1 '
Freiburg i. B., 1889.
138
INFLAMMATION.
ing very slight signs of movement thus arises and
grows gradually longer. These processes often meet
each other, and form loops which are at first solid
and afterwards become excavated with the forma-
tion of a vascular channel. Many observers, such as
Strieker, l Goloubew, 2 Klebs, 3 and Severini, 4 have
found that the endothelial walls of capillaries are to
a certain extent contractile, a still further proof of
a power of movement inherent in these cells. This
property must certainly play an important part in
the formation of stomata during inflammation, as
Klebs * thought, and as I imagined and stated in my
first papers on inflammation. 6 Arnold based his
Fig. 59. Passage of leucocytes through stigmata.
(After Arnold.)
original theory of stomata on the supposed pre-existence
of these orifices under normal conditions. This is not
the case however, since they are formed only at the
1 Moleschotfs Untersiichungen zur Naturlehre, vol. x.
2 Archil' f. mikroskop. Anat., 1868.
3 " Allgemeine Pathologic," 1889, vol. ii. p. 384.
4 " La contrattilita dei capillari," 1881.
5 See also Leydig, who in his pamphlet " Zelle und Gevvebe,"
1885, p. 17, has expressed the opinion that " in order to permit the
passage of the blood corpuscles, the minute pores may often dilate
into large orifices."
6 Biologischcs Centralblatt, 1883, p. 564.
LECTURE IX. 139
time of the inflammatory reaction, as he himself showed l
when he put forward his stigmata theory. (Fig. 59.)
Although Arnold does not allude to the contractility
of the endothelial cells, he admits that " the spaces be-
tween the cells alter according to the conditions of
tension and diffusion, so that the respective positions
of the endothelial cells may vary very greatly." 2 He
holds that the intercellular spaces are filled with a fluid
or viscous material. These pores between the endothelial
cells, which open to permit of the passage of the cor-
puscles and liquid parts of the blood, and close after
their passage, might with greater justice be compared
to the pores of the ectoderm in the Sponges, which
open and close on the passage of the corpuscles sus-
pended in the surrounding water. In both cases we
have to do with contractile cells whose movements may
be aided by the neighbouring cells (fusiform cells of the
Sponges and contractile cells of the endothelium of the
vessel wall).
The contractility of the endothelial cells may be
further shown by the fact that in some cases these cells
may leave the vessel wall and by means of their amoe-
boid movements wander into the lumen of the vessel.
This may be observed after injection of tubercle bacilli
into the circulation of rabbits. Some time after the
injection has been effected, it is found that the vessel
wall in some places is composed solely of the adven-
titia, the endothelial lining having entirely disappeared.
This is very frequently found to be the case in the
hepatic capillaries, where, as Eberth and others have
shown, the endothelium does not present the typical
1 Virchoiv's Archiv, 1875, vol. Ixii. p. 487.
* Ibid., 1876, vol. xlvi. p. 104.
I4O INFLAMMATION.
cell boundaries on treatment with nitrate of silver.
The endothelial cells of the hepatic capillaries become
readily detached from the outer coat, appearing in the
form of star-cells ' provided with a number of processes
which vary much in appearance. These cells, which
were first described by Kupffer, 1 were originally looked
upon as nervous elements on account of their shape
and thin elongated processes. It was soon ascertained
however that they belong to the endothelial tissue 2
and that they have the power of ingesting various
granules. They are frequently found, especially in
frogs, enclosing masses of reddish pigment, and they
readily take up granules of pigment, if these be injected
into the blood, as Ponfick 3 and others have shown.
Star-cells, in common with the other endothelial
cells of the vessels, have also the power of englobing
various pathogenic and saprophytic microbes which
have made their way into the blood. This is the case
with the leprosy bacilli, which have been found, some-
times in groups, in the endothelial cells of the blood-
vessels. They often occur in such large numbers that
they completely obscure the nucleus. This discovery
was made by Neisser 4 and Teuton, 6 who stated that
certain endothelial cells, when filled with bacilli, be-
come detached from the vessel-wall and are found free
in the lumen. Wyssbkowitch 6 has since found that
1 Archivf. mikroskop. Anat., vol. xii. p. 353.
2 Asch. " Ueber die Ablagerung von Fett und Pigment in den
Sternzellen der Leber." Bonn, 1884.
3 Virchow's Archiv, 1869, vol. xlviii. p. i.
4 Ibid., 1 88 1, vol. Ixxxiv.
6 Fortschritte der Medicin, 1 886, No. 2, p. 48. ( Virchow's Archiv,
1886, vol. civ. p. 381.)
6 Zeitschr. f. Hygiene, 1886, vol. i. p. i. See also the journal
Wratch, No. 44, 1891, p. 991.
LECTURE IX.
141
many microbes, when injected into the circulation of
rabbits, may be traced in the endothelial cells of the
hepatic capillaries. This may be observed some time
after the introduction of tubercle bacilli into the vein of
the rabbit's ear ; a large proportion of the bacilli are
found again in the endothelial cells, especially in the
liver. The same thing occurs in natural infections in
man. Thus after death
from malaria, the en-
dothelial cells of the
liver are found filled
with the characteristic
parasite of this disorder. 5
Perhaps the most strik-
ing instance is furnished
by pigeons that have
been inoculated with the
bacillus of swine septi-
caemia. Enormous mas-
ses of these microbes are
found filling nearly the
whole of the endothe-
lial Cells Of the blood- F|g 6o ._ Endothelia , cells enclosing the bacilli
vessels, especially in the [ g s n ne septica:mia ' from a hepa:ic vein of
liver. (Fig. 60.) As this bacillus, like that of leprosy, is
non-mobile, its presence within the protoplasm of the en-
dothelial cells can only be regarded as due to an active
ingestion on the part of the cells themselves.
I have laid weight on these details, since the contrac-
tility and the phagocj tic properties of the endothelial
cells are extremely important facts in the question of
inflammation. Other varieties of endothelial cells are
endowed with the same properties. Thus the lym-
142 INFLAMMATION.
phatic endothelial cells are still more often the seat
of leprosy bacilli than those of the blood-vessels.
Ranvier, 1 many years ago, showed that in the guinea-
pig the endothelial cells of the inflamed omentum were
capable of taking up foreign bodies (such as myelin or
vermilion) that had been injected into the peritoneal
cavity. In a recent study 2 of the inflammatory changes
in the omentum, he has been able to demonstrate the
contractility of the cells forming its endothelial cover-
ing. Moreover Ribbert, 3 in his memoir on the
inflammation and regeneration of lymphatic glands,
frequently lays stress on the contractility of the endo-
thelial cells of these organs.
The connective tissue cells certainly take some part
in inflammation, but this part is evidently much less
important or constant than was formerly thought, and
is far transcended by the part played by the leucocytes
and the endothelial cells of the blood-vessels. Further
we must remember that it is extremely difficult to make
sure of the co-operation of connective tissue cells in the
processes of inflammation. If they be studied in the
tails of tadpoles or of Urodelan larvae, the only changes
that we see in the fixed connective tissue cells are
alterations in their shape and their processes. These
changes last only a short time, and must therefore be
regarded as the immediate result of the lesion. These
facts prove that an absolutely typical inflammatory
reaction may occur without the fixed connective tissue
cells taking any important part in the process. These
cells, however, proliferate in an inflamed organ, and
1 " Le9ons sur le systeme nerveux," 1878, vol. i. p. 304.
s Comptes rendus de FAcad, des Sciences, April 20, 1891, p. 842.
3 Ziegler's Beitrage zur pathol. Anat., vol. vi. 1889, p. 205.
LECTURE IX. 143
so furnish a supply of new cells to replace those
killed by the original injury. Hence at a more or less
advanced stage of inflammation, karyokinetic figures
are to be seen in the connective tissue cells.
Ranvier l has recently called attention to a variety of
connective tissue cells corresponding to the " plasma
cells " of Waldeyer, which he designates as c/asma-
tocytes. (PI. II., Fig. 8.) These are very large
granular cells with processes and are formed from
leucocytes or migratory cells, which have increased in
size and lost their power of movement. During in-
flammation, according to Ranvier, the clasmatocytes
regain their former mobility, multiply rapidly and con-
tribute to the formation of pus-cells. On this view, the
clasmatocytes would form only a temporary constituent
element of the connective tissues, and would serve as a
source from which leucocytes might be drawn should
inflammation take place. These cells are present in
large numbers in the peritoneal tissue of tritons, frogs
and rabbits, but are not found in the larval stages of
amphibians.
Certain other cells of the connective tissue seem to
take some part in the process of inflammation. Thus
the basophile cells or " Mastzellen " of Ehrlich (PI.
III., Fig. 3) are often numerously represented in the
inflammatory products, although we are quite unable
to explain their presence under these circumstances.
Certain peculiarities in the reaction of these granules
to various stains seem to point to a possibility that
these cells may act as scavengers and clear away
the inflammatory products. Might we not indeed look
1 Comptes rendus de VAcad. des Sciences, April 27, 1891, p. 922.
144 INFLAMMATION.
upon the granules with which they are filled as the
excreta of the other cells ?
Having considered the principal cellular elements that
take part in the inflammatory reaction, we may now
inquire how they carry out their functions. I need not
insist at length on the fact that the migration of the
leucocytes through the vascular wall is due to their own
active movements. In spite of all Cohnheim's en-
deavours, in spite of the general desire to refer all vital
phenomena to mechanical causes, the view that the
emigration is effected by the amoeboid power of the
leucocytes has now found almost unanimous acceptance
(Thoma, von Recklinghausen, Lavdovsky and others).
This fact must strike anyone who compares the ease
with which leucocytes traverse the vascular wall, with
the purely passive diapedesis of the red blood-cor-
puscles. The latter often remain attached for an
indefinite time to the wall or are torn to pieces instead
of passing outside the vessels. Even Hering 1 who
originated the theory of the mechanical filtration of
leucocytes at the time of inflammation, admits the par-
ticipation of the amoeboid movements of these cells,
although not attributing any great importance to them.
According to him the blood must either be circulating
or be subjected to a certain pressure for the filtration of
leucocytes to take place. Now the complete emigra-
tion of leucocytes may readily be observed after entire
stoppage of the heart in tadpoles which have, as some-
times happens during an experiment, died from the
effects of the curare administered to them. (Fig. 61.)
1 Sitzungsbcrichte der k. Akad. der Wissensch. in Wien, 1868,
vol. Ivii., Part ii., p. 170.
LECTURE IX.
145
In the very act of the passage of the leucocytes
through the vessel-wall, a certain chemiotactic or
physiotactic influence is manifested. If we agree with
Massart and Bordet ! in considering this emigration as
a mere effect of tactile sensibility, we cannot explain
the fact that the leucocytes remain inside the vessels in
cases when these are sufficiently dilated but when there
may be in the neighbourhood substances which have a
repellent action on leucocytes. Some time ago Binz
found that no dia-
pedesis of white cor-
puscles took place
through the frog's
mesentery after this
had been moistened
with a solution of
quinine. As quinine
acts poisonously
Upon protoplasm, it F[ S- 6i.-Djapedesis of a leucocyte through the wall
* of a capillary, in which the blood is motionless.
was concluded that
it paralysed the leucocytes which were consequently
unable to pass through the vascular wall. The same
experiments were repeated by Disselhorst 2 who con-
firmed the fact that the diapedesis of the leucocytes
ceased, but was astonished to find that their movements
were not paralysed, for when removed from the vessels,
the leucocytes showed their wonted amoeboid activity.
In order to explain these facts, Disselhorst maintained
that the assistance of the vessel-wall was indispensable
for the passage of the leucocytes, and that the diapedesis
1 Journal published by the Soc. roy. d. Sciences medicates et
naturelles de Bruxelles, 1890, vol. v.
* Virchow's Archiv, 1888, vol. cxiii., p. 108.
146 INFLAMMATION.
was prevented by the action of the quinine upon the
wall. In his arguments, no account is taken of the
chemiotactic property of leucocytes, which was not at
that time an accepted fact. If this function be ad-
mitted, the effect of the quinine may be referred to a
negative chemiotaxis of the leucocytes, which, although
retaining their mobility, do not move towards the part
moistened with this substance. It follows that the
leucocytes are able, while still within the vessels, to
detect the presence of the quinine.
The same explanation may be applied to the cases
of infection by highly virulent microbes where the
leucocytes, in spite of the dilatation of the vessels, do not
cross the vascular wall. Thus in the most rapidly fatal
infectious diseases, such as chicken cholera, septicaemia
of guinea-pigs and pigeons, and many others, there is
scarcely any emigration towards the invaded spot. In
these cases no diapedesis takes place although the most
favourable conditions exist, i.e. considerable inflamma-
tion and hyperaemia accompanied with serous and
even haemorrhagic exsudation. This may be explained
by a negative chemiotaxis manifested by the intra-
vascular leucocytes. This view is moreover borne out
by the fact that the microbes of the above-mentioned
diseases are not englobed by the leucocytes when
brought into contact with them. The occurrence of
chemiotactic action within the vessels necessarily follows
from the leucocytosis produced by the absorption of
various substances formed by the bacteria.
Massart and Bordet succeeded in stopping diapedesis
by depressing the sensibility of the leucocytes with
paraldehyde and chloroform, although these drugs did
not paralyse their power of movement.
LECTURE IX. 147
It is highly probable that various other sensibilities,
besides those of touch and chemiotaxis, help in bring-
ing about the act of emigration. In lesions where the
necrosis of the cells is very slight and where conse-
quently the attraction due to chemiotaxis can only
be insignificant, we may probably regard the physical
change in the medium (difference of tensions, &c.) as
the chief factor in determining an attraction of the
leucocytes.
The diapedesis of the leucocytes is usually preceded
by the peripheral distribution of these cells in the dilated
vessel. This phenomenon, to which the name ' itio in
paries ' (Samuel) has been given, is generally considered
to be the most mechanical event in inflammation. It
was first sought to explain it by comparison of the
movements of the spherical white corpuscles with the
rapid motion of the flattened red blood discs ; but after-
wards the explanation suggested by Chklarewsky ' in
his account of a research carried out in Helmholtz's
laboratory, was adopted. This explanation is based on
the fact, discovered by Mach and Bondi, that insoluble
corpuscles when suspended in a fluid, increase the
density of the mixture. Now as the corpuscles circu-
late only in the axial portion of the vessel, while the
clear plasma flows along the sides, the density of the
latter must be less than that of the central portion. As
the leucocytes have a lower specific gravity than the
red blood- corpuscles, they are driven from the denser
axial current to the peripheral zone.
Although we must admit that the peripheral distribu-
tion of the leucocytes can only be due to a mechanical
action, since according to Massart and Bordet the
1 P finger's Archiv fur Physiologic, vol. i.
L 2
148 INFLAMMATION.
white corpuscles, after being anaesthetised by chloro-
form, still proceed towards the sides of the vessels, yet
we are unable to adopt the generally received explana-
tion of this phenomenon. It has been known since
Cohnheim's time and confirmed by Chklarewsky him-
self as well as by other observers that the peripheral
distribution, as well as emigration, goes on just as well
after the leucocytes have taken up vermilion granules.
As these leucocytes now contain a mercurial salt, they
must be not only heavier than they were before, but
also heavier than the red blood-corpuscles. Neverthe-
less the latter remain in the axial current while the
weighted leucocytes proceed towards the periphery.
The attraction of the leucocytes for the vascular wall
has been referred by Hering to their adhesiveness.
This theory has, however, found few supporters, for the
reason that leucocytes are not sticky and do not be-
come attached on account of their consistency but
solely by means of their amoeboid properties. Anyone
may be convinced of the fact that naked protoplasm
is not sticky by feeling the large plasmodium of the
Myxomycetes, such as that of the Spumaria alba.
If it were possible to admit that the sensibility of the
leucocytes in Massart and Bordet's experiments was
not completely abolished by the action of chloroform,
we might attribute their accumulation in the peripheral
zone of the dilated vessels to a remnant of sensibility by
which they could still appreciate the differences in the
external medium, and so would move towards the calmer
regions where they are free to extend their protoplasmic
processes.
The vascular wall, by virtue of the power of contrac-
tion possessed by its endothelial cells, may no doubt
LECTURE IX. 149
aid the passage of the leucocytes, although the latter
are quite capable of traversing non-contractile mem-
branes such as the epithelial tissues of vertebrates
(Stohr) or Ascidians. But the contractility of the
endothelial cells must have more influence in regu-
lating the passage of the red blood-corpuscles and the
liquid parts of the blood, especially in those cases where,
as in the infectious diseases mentioned above, the leuco-
cytes in consequence of negative chemiotaxis remain
in the blood, while the plasma and blood-discs exsude
through the vascular wall.
Although the leucocytes may emigrate from the
venules even when these are normal or only slightly
dilated, (as Recklinghausen first observed in tadpoles
under normal conditions,) it is nevertheless undeniable
that the dilatation of the vessels accompanied by slow-
ing of the blood-stream offers a very favourable condi-
tion for the occurrence of diapedesis. This dilatation,
which is so constant in the inflammation of vertebrates,
is a very complicated phenomenon in which several
factors take part. Cohnheim, 1 who ascribed it exclu-
sively to the action of the vascular wall itself, sought to
prove his point by the following experiment. After
pinning out the tongue of a frog on a disc of cork, he
cut all the tissues connecting it with the body, with the
exception of the artery and vein. In this way he con-
sidered that he had eliminated any possible nervous
influences. He then applied some irritant to the
tongue, and found that an inflammatory reaction was
produced as usual. Cohnheim would not admit the
possibility of local nervous mechanisms in the vascular
1 " Gesammelte Abhandlungen." 1885, p. 423.
1 50 INFLAMMATION.
wall, although it is very probable that such mechanisms
do play a part in these phenomena.
A considerable importance in the production of in-
flammatory hyperaemia is generally ascribed to the
state of tension in the tissues, which is supposed to
react on the capillaries. This idea, started by Fluss,
was afterwards supported by Landerer, 1 who undertook
a series of experiments to show that the tension in the
tissues during inflammation is much lower than under
normal conditions. The capillaries surrounded by
these inflamed tissues are unable to withstand the
pressure of the blood and therefore dilate. The imme-
diate effect of this dilatation is an acceleration of the
movement of the blood, which rushes in to fill the
widened capillaries. But after a certain lapse of time,
as the surrounding tissues lose their elasticity more and
more, the blood-stream slows, thus producing the con-
dition of the circulation typical of inflammation.
This theory also suffers from the disadvantage that
it takes no account of the part played by nervous in-
fluence, which has been so much studied of late years.
Samuel 2 has in the following experiment strikingly
shown the importance of this influence. After cutting
the sympathetic nerve in a rabbit on one side, he
divided both auricular nerves on the other side of the
same animal. Paralysis of the vaso-motors at first pro-
duced congestion of the ear on the side of the divided
sympathetic and, as an indirect effect, a certain degree
of anaemia in the opposite ear. This anaemia increased
greatly after section of the sensory nerves, on account
1 " Ueber die Gewebsspannung," 1884, and "Zur Lehrevon der
Entziindung." Leipzig, 1885.
2 Virckovfs Afckiv, 1890, vol. cxxi., p. 396.
LECTURE IX. 151
of the impossibility of reflex action by the uninjured
sympathetic. If, however, inflammation was produced
by treating both ears with hot water (at 54), totally dif-
ferent results were obtained. The ear on the side of
the divided sympathetic became very hyperaemic and
highly inflamed. In the other ear, no hyperaemia
occurred, and inflammation was replaced by a stasis
ending in gangrene. This experiment proves the
influence of the nervous system on inflammatory re-
action and at the same time the beneficial effects of the
latter. On the side where the inflammation is ex-
aggerated in consequence of the paralysis of the vaso-
motor nerves, the ear is speedily restored to a normal
condition, whereas the morbid phenomena are excep-
tionally severe on the anaemic side, where, owing to the
division of the sensory nerves, the processes of in-
flammation are unable to take their usual course.
In the rabbits in which Samuel divided the sensory
nerves (auricularis major and minor) on one side only,
the results were much less serious. If the ears of these
rabbits were dipped into water at 54, in adult animals
the inflammation followed the normal course, and it
was only in young animals which had been treated
in this manner that he observed a retardation of the
inflammatory reaction and of the subsequent healing
process.
Similar experiments have been made by Roger. 1
But instead of producing inflammation with hot water,
he made use of the cultures of the streptococcus of
erysipelas. He first divided the sensory nerves of one
ear of a rabbit and then inoculated both ears with these
bacteria. In the ear of which he had divided the
1 Comptes rendus de la Soc. de Biologie, 1890, No. 34, p. 646.
152 INFLAMMATION.
auricular nerves, the erysipelas lasted a long time, and
the healing processes, which were very slow, were
accompanied by some mutilation of the organ, whereas
in the normal ear of the opposite side the erysipelas ran
its usual course.
The contrary result was obtained by Roger l on
inoculating rabbits with erysipelas, after dividing the
sympathetic instead of the sensory nerves on one side.
The paralysis of the vaso-motors thus produced gave rise
to a condition of hyperaemia, which exercised a favour-
able influence on the course of the erysipelas compared
with the other ear where the sympathetic was intact.
Inflammation began much earlier and the ear healed
more rapidly on the side on which the sympathetic had
been divided than on the other, although here also a
cure finally resulted.
These experiments confirmed those of several other
authorities, as Snellen, K. Danilewsky and others, who
had observed an improvement in the course taken by
the inflammation after division of the cervical sym-
pathetic nerve. Since section of this nerve on one side
also affects the ear of the opposite side by rendering it
anaemic, Samuel modified his experiment in the follow-
ing way. Instead of setting up inflammation in both
ears of the same rabbit, he made use of two different
rabbits, in one of which he divided the sympathetic on
one side, while the other animal was left uninjured.
He produced inflammation in the operated rabbit by
the application of water at 54 to the ear on the side of
the section, and in one ear of the normal rabbit by the
same means. He then found that the inflammation
1 Comptes rendus de la Soc. de Biologie, 1890, No. 16, p. 222.
LECTURE IX. 153
ran a milder course in the control animal than in the
operated rabbit.
All these experiments prove that the nervous system
has a certain influence on inflammation, but at the same
time shows that the influence only serves to accelerate
or retard its course. While duly acknowledging that
it plays this part, we must be careful not to exaggerate
its importance, as by so doing we may fail to recognise
the true relations of the different factors which co-
operate to produce the inflammatory reaction.
It has been shown by Charrin and Gley l that reflex
dilatation of the vessels can be prevented by injection
of 20 c.c. of the soluble products of the bacillus pyocy-
aneus into the circulation of a rabbit. After this injec-
tion, these authors observed that " the vasodilatation
thus produced develops more slowly, besides being less
marked and of shorter duration," (p. 734) than under
normal circumstances. In a later note ' z they proved
that this inhibitory influence on the vascular dilatation
is due exclusively to the volatile products contained in
the cultures and is not shared by the non-volatile con-
stituents, whether soluble or not in alcohol. These
facts they refer to a diminution of the excitability of
the vasodilator mechanism under the influence of these
volatile products. According to them the secretions of
the microbes diminish dilatation, hinder diapedesis and
in this way favour the course of the infection. They
seek to apply these results to the explanation of
immunity ; thus vaccination would act by strengthening
the part of the nervous system which presides over the
dilatation of the blood-vessels and over diapedesis.
1 Archives de Physiologic, No. 4, 1890, p. 724.
2 Ibid., No. i, 1891, p. 146.
154
INFLAMMATION.
Charrm and Gamalei'a l have likewise prevented the
dilatation of the blood-vessels by the injection of
vaccines or of the products of the micro-organism of
vibrionian septicaemia, as well as by simple injections
of 5-10 per cent, solutions of salt.
We cannot suppose that in the natural course of an
infective disorder there is a sudden entry into the blood
of 20 c.c. of bacterial products. In order then to
come to a decision on the subject, we must experi-
ment on animals which have been inoculated with the
microbes in question. If we introduce some bacillus
pyocaneus from a culture on agar-agar under the skin
of two rabbits, one of which has not been treated, while
the other has been vaccinated against the malady pro-
duced by this bacillus, the inflammation produced is
much more marked in the first rabbit than in the
vaccinated animal. In the former the bacterial pro-
ducts prevent neither the vascular dilatation nor the
increased local temperature, nor the serous exsudation
at the point inoculated ; and yet the diapedesis is less
than in the vaccinated rabbit, in which on the other
hand the vasodilatation and increase of temperature
are much less pronounced. The difference is still
more striking if we compare the results of injecting
the vibrio of septicaemia ( Vibrio Metchnikowii} under
the skin of the ear in two guinea-pigs, one of which is
susceptible, while the other has been rendered refrac-
tory by means of vaccinations. Here we find that the
reddening, heat and tumefaction are much more marked
in the susceptible than in the immune animal, whereas
diapedesis which is well marked in the vaccinated
animal scarcely occurs at all in the other.
1 Centralblatt f. allgemeine Pathologie, Nos. 18, 19, 1890, vol. i.,
p. 588.
LECTURE IX. 155
These facts show in the first place that, contrary to
the conclusions of Charrin and Gley, the microbes in
question do not hinder the vascular dilatation in sus-
ceptible animals ; and, in the second place, that dia-
pedesis may be almost or entirely absent in spite of
considerable dilatation of the blood-vessels. We must
conclude therefore that the most important factor in
the production of diapedesis is the sensibility of the
leucocytes themselves. If this sensibility be positive,
the leucocytes emigrate, in spite of the insignificant
dilatation of the vessels ; if it be negative, diapedesis is
not produced, however dilated the blood-vessels may be.
This fact may be easily shown by using for inoculation
a microbe exercising considerable positive chemiotaxis
on the leucocytes. If two guinea-pigs be inoculated
under the skin of the ear, one with a small amount of
tubercle bacilli and the other with the vibrio Metch-
nikowii, in the first animal the dilatation will be slight
and the diapedesis very considerable, while in the latter
the dilatation will be marked but will be accompanied
by little or no diapedesis.
The following facts may also serve as indirect argu-
ments against Charrin's and Gley's interpretation of
their results. In the most acute infections, in which
diapedesis is nearly or entirely absent, the presence of
leucocytes does not in any way hinder the existence and
multiplication of the bacteria, since in consequence of a
negative chemiotaxis the leucocytes do not englobe
these microbes. Under these circumstances it would be
quite superfluous to hinder diapedesis. If then, instead
of introducing some of these malignant bacteria (such
as the bacillus of chicken cholera in birds and rabbits,
or the vibrio of avian septicaemia in guinea-pigs and
I5 6 INFLAMMATION.
pigeons) into the subcutaneous tissue, we introduce them
directly into the blood, the result is exactly the same.
The animals die in a very short time without the
occurrence of any phagocytosis ; and it makes no
difference to the issue whether, as in this case, the bac-
teria are in close proximity to the leucocytes in the
blood and spleen, or whether, as after subcutaneous in-
oculation, they are at some distance from these cells,
which remain in the interior of the blood-vessels.
We must therefore look upon the sensibflity of the
leucocytes as the most important factor in inflammatory
diseases, not forgetting however the part played in the
inflammatory reaction in vertebrates by the sensibility
of the endothelial cells, as well as by that of the nervous
and other elements. 1
1 See my lecture on " Immunity " in the British Medical
Journal of January 3ist, 1891. After I had written this chapter,
Massart and Bordet put forward the same ideas in a study of the
chemiotaxis of leucocytes in infectious disorders (Annales de
rinstitut Pasteur, 1891, p. 417). They concluded from these ex-
periments that the absence of diapedesis in certain infectious
disorders is due not to a paralysis of the vasomotor centres, but
rather to a negative chemiotaxis of the leucocytes. In consequence
of this paper, a controversy has arisen between Charrin and Gley
on the one side and Massart and Bordet on the other (Comptes
rendus de la Soc. de Biologic, 1891, pp. 703-710. See also
Bouchard, Comptes rendus de FAcad. des Sc., 1891, pp. 524-529).
Without entering into the details of this discussion, we may state
generally that it is not a vasomotor paralysis but a negative
chemiotaxis which is the real cause of the absence of leucocytes at
the seat of the lesion in certain severe bacterial infections. Hertwig
(" Physiologische Grundlage der Tuberculinwirkung." Jena, 1891)
and Buchner (Munchener meditinische Wochenschrift, 1891) have
also pronounced in favour of this view.
LECTURE X.
Chronic inflammations Tuberculosis as a type of a chronic in-
flammation Phagocytic nature of tuberculous cells Destruc-
tion of tubercle bacilli by phagocytes Power of resistance of
Meriones to tubercle Leprosy.
HAVING passed in review the principal elements in-
volved in inflammation in the .vertebrata, it remains to
inquire whether the part played by each is invariably
the same in all cases of inflammatory reaction. In
acute inflammation we find as a general rule vascular
dilatation accompanied by an active condition of the
endothelium of the vessel-walls and an exsudation with
diapedesis, that is to say, three events which concur in
producing a considerable afflux of leucocytes towards
the injured spot. Is the order of phenomena the same
in the chronic inflammations ? We often meet with
the idea that whereas in acute inflammation vascular
changes accompanied by diapedesis play the most
important part, in chronic inflammation the greatest
significance must be attributed to local phenomena that
take place in the tissues without the co-operation of the
cells of the blood or blood-vessels.
Chronic, like acute inflammations, are due to various
causes, which may be chemical, physical, or, as is more
usually the case, biological in nature. They may be
158 INFLAMMATION.
brought about by the slow action of any injurious sub-
stance, as lead, phosphorus or alcohol, or by the pro-
longed action of heat or other harmful physical means ;
but they are more frequently the immediate effect of
microbes and their poisonous products.
We may take as an instance the chronic inflamma-
tion which results in the production of tubercles in
bacillary tuberculosis or in some other granuloma. We
choose this type, not only on account of its great im-
portance in pathology, but also because the fixed ele-
ments of the tissues have been regarded as the chief
factors which contribute to its formation. According to
Baumgarten's theory, 1 which is accepted by most patho-
logists, the phagocytic cells in general and the leuco-
cytes in particular take little or no part in the formation
of the tubercle itself. This he considers to be produced
by a proliferation of the local tissue cells, excited by the
presence of the tubercle bacillus in their neighbourhood.
Thus, according to this view, pulmonary tubercle is de-
veloped at the expense of the epithelial cells of the
alveoli ; hepatic tubercle at the expense of the
liver-cells and the epithelial cells of the bile-ducts ;
renal tubercle by a proliferation of the epithelial cells of
the urinary tubules, &c. The connective tissue also
contributes to the formation of tubercle by means of its
fixed cells and endothelial cells. The leucocytes take
no part in the formation of the tubercle until a later
period and hence must be regarded as of merely
secondary importance. According to this theory,
tuberculosis would not be a true inflammation, other-
wise we must admit that a chronic inflammation may
1 " Tuberkel und Tuberkulose." Berlin, 1885. Reprinted from
the Zeitschrift f. klin. Median.
LECTURE X. 159
be developed almost entirely independently of the
phagocytes or of the leucocytes.
The formation of tubercle may be accelerated by in-
jecting bacilli of avian tuberculosis into the veins of
rabbits, which, as is well known, are very susceptible to
this disorder. At the end of a few days, microscopic
tubercles are developed which may serve as a type of
this class of neoplasm. If we examine the liver, we shall
see that the epithelioid and giant cells of the tubercles
are formed solely at the expense of the phagocytic ele-
ments, that is to say, of the large mononuclear leuco-
cytes and of Kupffer's star-cells, derived from the
vascular endothelium. No hepatic or epithelial cells
ever contribute to the production of tubercle. It is true
that their nuclei may occasionally be seen about to
undergo karyokinetic division ; but this proliferation
has no direct connection with the formation of tubercle,
and merely serves for the regeneration of the hepatic
cells themselves.
Hepatic tubercle is the result of an accumulation, not
a multiplication of the mesodermic phagocytes, which
collect in masses and make up the whole of the original
tubercle. These phagocytes contain bacilli which have
been englobed by the amoeboid cells. Several of the
epithelioid cells fuse together to form giant cells, while the
tubercular elements are reinforced by a large number of
lymphocytes and mononuclear leucocytes. Although
the former are not yet leucocytes, they soon become
converted into epithelioid cells, which represent one
variety of leucocytes.
Hepatic tubercle is therefore not derived from various
sources, as Baumgarten maintains, but is produced by
the amoeboid and phagocytic cells of the mesoderm.
i6o
INFLAMMATION.
The mode in which pulmonary tubercle is developed
confirms this view. This variety of tubercle is formed
by the endothelial cells of the blood-vessels together
with the leucocytes, and is the result not of the proli-
feration but of the accumulation of these cells, which is
effected by means of their amoeboid movements. The
cells of these tubercles likewise show their phagocytic
properties by englobing the tubercle bacilli. In cases
where 'dust-cells' (' Staubzellen ') contribute to the
formation of tubercles, we have again to do with phago-
cytic elements derived from mononuclear leucocytes. 1
Tubercles of the spleen and of the lymphatic glands
are likewise formed by an agglomeration of the larger
phagocytes of these organs phagocytes possessing a
single large nucleus. In guinea-pigs and marmots
(Spermophilus] a study of the neoplasms produced by
the bacillus of avian or human tuberculosis further con-
firms the statement that tubercle is composed of a collec-
tion of phagocytes mesodermic in origin, which move
towards the spot where the bacilli are situated and englobe
them. The phagocytes retain their condition of epithe-
lioid cells, or are transformed into giant cells. The
latter can also be formed in another way, although in
any case the final result is a large mass of protoplasm
enclosing a number of nuclei. In some animals, as the
marmots, the nuclei proliferate by a process of budding ;
more frequently however the multiple nuclei are
derived from a number of cells which have fused to
form a plasmodium. It is possible that the nuclei may
1 See Tchistowitch, Annales de Hnstitut Pasteur, 1889, p. 337.
Afanasieff, in an investigation carried out in my laboratory on the
development of pulmonary tubercle in the rabbit after inoculation
of tubercle bacilli from man, proved that here also the tubercle is a
purely phagocytic product.
LECTURE X. l6l
also multiply by karyokinesis, but this has never been
demonstrated with certainty.
It is a well established fact that leucocytes take part
in the formation of tubercle, but these leucocytes belong
to the mononuclear variety. The polynuclear cells
englobe the tubercle bacilli readily but perish after a
short time, and then with the microbes they contain, are
eaten up by various mononuclear phagocytes which"
may be classed together under the term of macrophages.
These latter cells have a much greater power of resis-
tance, and in some cases are even capable of destroying
the tubercle bacilli. Thus after inoculating marmots
with avian or human tubercle I have observed a very
characteristic degeneration of the bacilli in the interior
of the epithelioid and giant cells of these animals, which
are but little susceptible to tuberculosis. 1 The bacilli,
evidently under the influence of these cells, swell up
and gradually lose their power of retaining colouring
reagents. Sometimes the peripheral part but more
often the central part of the bacillus is the first to lose
its coloration. Later on the bacillus becomes converted
into a yellowish sausage-shaped body, enclosing a very
delicate canal, and the altered microbes collect to form
a conspicuous brownish mass, having the characteristic
aspect of a fragment of amber. None of these changes
1 See my article in Virchow's Archiv, 1888, vol. cxiii., p. 63.
The facts brought forward in this paper concerning tubercu-
losis in marmots refer to the bacillus of avian tuberculosis.
Later experiments have shown me that this animal is more suscep-
tible to the bacilli of human tuberculosis, although, even in this
case, a certain number of bacilli are destroyed in the cells and
converted into yellow particles. At the Congress in London,
Bardach brought forward facts showing the high degree of suscep-
tibility possessed by the marmots for the virus of human tubercu-
losis, which he made use of in his experiments.
M
162 INFLAMMATION.
are ever observed in artificial cultivations, even though
a number of dead bacilli are present, nor indeed any-
where outside the tubercular cells. They must be there-
fore regarded as resulting from the phagocytic action
of these cells, and recall the degenerative phenomena
which have been described in an earlier chapter as taking
place in cysts of the Gregarinae and the larvae of the
Nematoda, when surrounded by the phagocytes of
Lumbricus. In all these cases we have to do with
abnormal secretions poured out by the parasites in order
to resist the influence of the phagocytes which enclose
or surround them.
Bacilli have also been ascertained to undergo changes
similar to those just described, within the giant cells of
rabbits, and very occasionally within those of guinea-
pigs. On the other hand I have never been able to
detect bacilli which have been destroyed by these means
either in the tuberculosis of Bovidae or of man. And
yet the resistance offered by the organism in such cases
is often of a very marked character. Calcareous de-
generation of tubercle has long been recognised as a
means by which tuberculosis in human beings may be
cured. In order to give a better idea of this reactive
phenomenon, I will describe the manner in which the
Algerian rat (Meriones] succeeds in resisting the action of
the tubercle bacillus. This rodent, although not entirely
immune against tuberculosis, receives it in a milder
form than many of the allied species. If inoculated
under the skin or even in the eye with a culture of the
bacilli of human tuberculosis, this animal withstands
the effects of the disease for many months.
If an Algerian rat that has been inoculated six or
eight months previously, be killed, a large number of
LECTURE X.
163
tubercles are found in the abdominal organs, the lungs
and the lymphatic glands. These tubercles, however,
in the majority of cases, do not present any necrosis or
caseation. The tubercular tissue composed of living
cells encloses bacilli, the majority of which have under-
gone a very remarkable degeneration, which deserves
our further consideration. In this animal the main
struggle goes on in the spleen, which is studded with
small tubercles consisting of non-necrosed epithelioid
and giant cells. The
former cells enclose a
small number of ordi-
nary tubercle bacilli,
while the giant cells
contain characteristic
calcareous bodies. (Fig.
62.) When examined
under the microscope
they are seen to be high-
ly refracting and in the
majority of cases to have
the shape of a figure
of eight, although some-
times they are simply rounded or irregular in form.
Under the influence of acids, the lime salt (phosphate of
lime) may be dissolved out, leaving a skeleton consisting
of a larger or smaller number of fine concentric layers.
(Fig. 63.)
These calcareous bodies have a strong resemblance
to the formations described by Schuppel l as occurring
Fig. 62. Tubercle from Meriones enclosing
a calcareous body.
1 " Untersuchungen iiber Lymphdriisen-Tuberkulose.' :
gen, 1871, p. 104 and PI. I., Fig. 3, 4.
M 2
Tiibin-
1 64
INFLAMMATION.
in scrofulous glands, and observed by several authors l
in many cases of tubercle of the lymphatic glands in
man. (Figs. 64, 65.) But whereas the origin of the
striated calcareous bodies in man is still quite obscure,
it can be demonstrated in Meriones with the utmost
facility. The examination of cover-glass specimens or
sections doubly stained by the methods of Gram or
Ziehl, shows at once that these bodies are the result of
a degeneration of the tubercle bacilli in the interior of
the giant cells. In the early stages, the bacilli stain in
the usual manner, and do not present any noticeable
Fig. 63. The same cal-
careous body treated
with a dilute acid.
Fig. 64. Tubercle contain-
ing a calcareous body.
From a human scrofulous
gland. (After Schiippel.)
Fig. 65. Calcareous
body fram a mesen-
teric gland of man.
(After Ziegler.)
features. But by the side of these, we see other giant
cells (Tab. III., Fig. 4), in which the bacilli (a) are
covered with a fairly thick layer of an amorphous,
colourless substance (). This secretion becomes more
and more abundant so that the bacilli are finally sur-
rounded with a number of concentric layers. In many
cases these microbes may be stained with gentian
violet or fuchsin in the usual manner, but more fre-
quently they lose this power and take up the comple-
mentary stain. (Tab. III., Fig. 5.) Sometimes in the
centre of a calcareous body, we meet with a bacillus
divided into two, half of which still retains the primary
1 Ziegler, " Lehrbuch der patholog. Anat.," 6th Edition, 1890,
vol. ii., p. 98, Fig. 50.
LECTURE X. 165
coloration while the other has already lost it. (Tab.
III., Fig. 6.) Thus passing through a series of inter-
mediate changes, the bacillus arrives at a condition
when it will no longer take the stain, although its
contour is still well-marked. (Tab. III., Fig. 7.)
Later on, however, they are not to be distinguished
by their coloration from the surrounding substance,
(Tab. III., Fig. 8) and finally disappear completely.
(Tab. III., Fig. 9.) In the last stage, which is most
frequently met with, we see nothing but the stratified
calcareous bodies.
The concentric membranes, which are left after the
removal of the phosphate of lime by means of dilute
acids, consist of a substance identical with that which
constitutes the envelope of the tubercle bacillus. Like
the latter, they are dissolved by concentrated acids,
they are not affected by alkalies, and do not give a red
colour with Millon's reagent. These facts must lead us
to the conclusion that the concentric layers are secre-
tions of the tubercle bacilli and are exactly analogous
to the stratified cuticle which, as we have seen, becomes
formed round the Gregarinae and Nematoda when
attacked by the phagocytes of Lumbricus (p. 69).
Thus in Meriones the bacillus defends itself against the
giant cell in which it is lying by the secretion of a series
of envelopes. On the other hand it is very probable
that the phosphate of lime is laid down by the giant
cell itself in its reaction against the bacillus.
The struggle between these two living bodies the
tubercle bacillus and the giant cell of Meriones is
thus carried on by means of secretions. The bacillus
defends itself by the secretion of cuticular membranes,
and probably also by the production of toxines, while
1 66 INFLAMMATION.
the giant cell secretes a calcareous deposit by means of
which it walls in the bacillus, and usually succeeds in kill-
ing it. The giant cell probably also produces digestive
fluids which aid it in attacking and digesting the bacillus.
For the deposition of phosphate of lime to take place
in the interior of the giant cell an alkaline medium is
required. The alkaline reaction of the contents of the
giant cell and of the calcareous body formed from the
bacillus may be shown by various reagents such as the
sulpho-acid of alizarin, which is a very delicate test for
the presence of alkalies and gives these structures a
pronounced violet colour. Their intense coloration
with haematoxylin points to the same conclusion (Tab.
III., Fig. 8).
The exact conditions of this struggle between the
microbe and the phagocyte are highly interesting and
important and will form the subject of a separate
experimental investigation. Here I will only lay stress
on the conclusion that the giant cells, so characteristic
of tubercle, really represent a special form of phago-
cytes, which are particularly energetic in the conflict
with the microbes. This is moreover confirmed by the
discovery of Soudakevvitch, 1 that the giant cells of lupus
are able to digest the elastic fibres of the skin. To
attack substances so resistant as tubercle bacilli and
elastic fibres requires very active power of digestion.
In this respect the giant cells of tubercle may be com-
pared to the giant cells known as osteoclasts, which
effect the absorption of bone.
Weigert, 2 whose view on this point is accepted by
1 Virchoirfs Archh', 1889, vol. cxv., p. 264.
* Deutsche med.Wochenschr., \%%^^^ Fortschritte der Medicin,
1888, p. 809.
LECTURE X. 167
Koch L and other pathologists, considers that the giant
cell of tubercle represents a condition of partial necrosis,
by which the division of the protoplasm is prevented.
Against this theory, besides the above-mentioned fact
of the remarkable power of resistance possessed by
these cells, we may cite their property of dividing into
smaller cells. It is not surprising that this division
should be unaccompanied by karyokinetic phenomena,
since here the nuclei are multiple to start with, and all
that is still necessary is the division of the protoplasmic
body of the cell.
If we accept the opinion put forward by various
authors and lately revived by Chun 2 that fragmentation
of the nucleus, by enlarging its surface, may serve to
heighten the activity of the intracellular processes of
nutrition, it is easy to understand the teleological reason
for the increase in the ' number of nuclei in the cells
which have to digest bodies so resistant as those in
question. We must not forget that in many inverte-
brate animals plasmodia are often formed during the
process of intracellular digestion, these fused agglomera-
tions of cells being very commonly met with in the
phagocytic reaction of Sponges, Medusae, Echinoderms,
Molluscs, Daphniae, &c.
The cells of tubercle, which serve the organism in its
reaction against the microbes, are often destroyed by
the action of the bacilli, and then form the necrosed
caseous masses so characteristic of this disorder. This
degeneration is probably due in some degree to the
fact that the tubercles do not contain any blood vessels,
so that the activity and vitality of the phagocytes suffer
1 Deutsch. med. Woch., 1891, No. 3, p. 102.
* Physik.-okon. Gesellschaft zu Konigsberg. April 3rd, 1890.
1 68 INFLAMMATION.
from lack of nourishment. Koch has found that if the
inflammatory process in the tubercles be increased by
injection of tuberculin, a striking improvement may be
produced in the condition of guinea-pigs, and many
experimenters have observed a temporary amelioration
in man, alter the same procedure. Probably in this
case the cells of the tubercles are better nourished in
consequence of the inflammatory hypersemia that is
set up, so that they become more active and better
fitted to resist the action of the bacilli.
The type of chronic inflammation that we have
chosen has shown us that the essence of the process
consists in a reaction of mesodermic phagocytes
against the offending agent. Although the macro-
phages are more conspicuous than the leucocytic ele-
ments, yet we must remember that among the macro-
phages of tubercles a large number of mononuclear
leucocytes also occur ; and Ehrlich l has shown that
the latter elements are markedly increased in the
blood of phthisical patients.
Another chronic disorder, leprosy, which is analogous
in many respects to tuberculosis, must also be regarded
as a struggle between the mesodermic phagocytes and
the specific bacillus a struggle which lasts for many
years. Although the origin of the macrophages is not
yet known with certainty, it is probable that they are
derived from the endothelial and connective tissue cells,
as well as from the mononuclear leucocytes. These
cells unite to form granulomata, and destroy the bacilli
that they englobe, thus acting as phagocytes.
1 Charite Annalen, 1887, vol. xii., and " Farbenanaiytische
Untersuchungen,'' 1891, vol. i., p. 124.
LECTURE X. 169
The older researches of Neisser ' and Kebner, 2 as well
as the more recent ones of Soudakewitch 3 and Saw-
tchenko 4 leave no doubt as to the general characters of
the cellular pathology of leprosy. Having had several
opportunities of studying leprosy of the skin and of the
liver (among others in the preparations of Muskatbluth
made in my former laboratory at Odessa), I am able to
confirm the fact that in these cases the leprous cells
possess all the characteristics of mesodermic phagocytes.
In the liver the cells which englobe the leprosy bacilli
are chiefly of the endothelial type, with an extra-
ordinary number of vacuoles in their protoplasm.
From the point of view of the phagocytic theory, this
vacuolar ' degeneration ' should rather be regarded
as due to an abundant secretion of digestive fluids,
analogous to that observed in the Protozoa while intra-
cellular digestion is going on. In the interior of
leprous cells, especially those of the skin, we often find
enormous numbers of dead bacilli, showing the thera-
peutic action of these cells. The fact that Boinet and
Borel* have not been able to confirm this function of
the cells shows either that they had to do with special
cases, or what is more probable, that their observations
were incomplete.
Among the tubercular diseases one that has been
recently studied by Ebstein and Nicolaier 6 has a
special interest for us, viz. : the tubercular affection
1 Virchoitfs Archiv^ 1881, vol. Ixxxiv., p. 520.
2 Ibid., 1882, vol. Ixxxviii., p. 299.
3 Ziegler's Beitrdge zur pathol. Anat., vol. ii., Pt. i.
4 Ibid., vol. ix. p. 241.
& Comptes rendus de la Soc. de J3iol., 1 890, p. 38.
6 Vircho-w's Archil', 1889, vol. cxviii., p. 432, pi. xiii., xiv.
I/O INFLAMMATION.
produced by Nematoda in the kidneys of dogs and
the lungs of cats. In these cases the larvae become
surrounded by veritable tubercles, composed in the
dog of epithelioid cells alone, and in the cat of these
together with giant cells. In the dog each tubercle
contains a living larva enclosed by masses of tubercular
cells. Here then we have an undoubted instance of
an aggregation of leucocytes around a living animal,
recalling forcibly the collection of phagocytes in
Lumbricus round the living larvae of Rhabditis, as de-
scribed in the fifth lecture. It demonstrates once more
the fact that an accumulation of leucocytes may be
provoked by the presence of living organisms, and
does not require the presence of dead substances, or
substances derived from the dead bodies of the para-
sites. Unfortunately, however, these cases ofzoopara-
sitic tuberculosis have not yet been sufficiently studied
to give us any accurate knowledge as to the origin or
development of the cellular elements composing the
tubercles.
As Brault l has pointed out, there is a striking
analogy between the chronic inflammation brought
about by the tubercle bacillus and the processes that
give rise to the disease known by the name of liyper-
trophic cirrhosis. In both cases we have a reaction of
the connective tissues, or rather of the mesodermic cells
in general, a reaction which extends over a prolonged
period. Even if we regard it as an established fact that
the cirrhoses are produced directly by chemical poisons
such as lead, alcohol and others, the analogy of the
pathological processes in the two instances is by no
means destroyed. In the chronic infectious inflamma-
1 Archives generates de medecine % 1888, p. 47.
LECTURE X. i;i
tions the phagocytes collect around the parasites,
whereas in the cirrhoses they attack the tiss ue-cells that
have been enfeebled under the action of the poisons.
The final results then in the two cases may be quite
different, since in one the phagocytes destroy the
microbes, while in the other they annihilate the cells
belonging to the organism itself. In this respect the
chronic inflammations are closely allied to the phe-
nomena of atrophy, which however are scarcely within
the range of these lectures.
LECTURE XI.
Serous inflammations Two classes of these inflammations
Bactericidal power of the humours and the serous exsudations
Antitoxic property of serum and the serous inflammations.
IN the chronic, as in the acute inflammations, whether
purulent, fibrinous or catarrhal, we find that a pro-
nounced phagocytic action forms the key note of the
whole process. In the serous inflammations, however,
the number of leucocytes present is too small to permit
of our ascribing any great significance to their presence.
Up to the present our knowledge of these serous in-
flammations is very incomplete. So far as we can
judge from experiments on animals, they may assume
various forms.
In the first place there are certain inflammations
which acquire the character of l serous ' in consequence
of a negative chemiotactic influence on the leucocytes,
which do not emigrate or move towards the seat of
lesion, and therefore do not take any part in the forma-
tion of the exsudation. The walls of the vessels,
however, react as usual, thus permitting the passage
through their pores of the fluid forming the serous ex-
sudation, which is almost entirely devoid of leucocytes.
The exsudation often contains numerous pathogenic
microbes which multiply rapidly in it without let or
LECTURE XL 1/3
hindrance. Deprived in this way of any phagocytic
protection, the animal organism speedily becomes a
prey to the microbes. In this class of serous inflam-
mations we may include the most acute and fatal
diseases, such as the vibrionic septicaemia of birds and
guinea-pigs, anthrax in the smaller rodents as mice and
guinea-pigs, and some other infectious disorders. In
the form of septicaemia just mentioned, the vibrio
swarms freely in the serous exsudation with its almost
total absence of leucocytes. In the serous exsudation
of animals infected with anthrax, the bacilli are only
present in small numbers, although the fluid collects
at or in the immediate neighbourhood of the infected
spots.
In a second group of serous inflammations of microbic
origin, the exsudation is formed in regions, which may
be at some distance from the deposit of microbes, and
does not contain any bacteria at all. Roux and Yersin '
have shown that the exsudation produced in the serous
pleurisy which often accompanies diphtheria in guinea-
pigs does not contain any microbes, which remain
localised at the seat of inoculation. In ten cases of
serous inflammations accompanying phlegmonous ery-
sipelas or other suppurative maladies, the liquid con-
tained in the inflammatory oedema was found by
Zimmermann 2 to be absolutely free from microbes.
The question at once suggests itself whether this trans-
sudation of fluid really represents a phenomenon of
reaction on the part of the organism, and if so, what
use it is to the infected organism.
In examining this point we naturally think first of a
1 Annales de Vlnstitut Pasteur, 1888, p. 635.
2 Munchener medicinisclie Wochenschrift, 1889, No. 9, p. 141.
174 INFLAMMATION.
possible microbicide action of the transsuded liquid
which would rid the organism of its aggressors. We
find, however, that all kinds of bacteria can live and
flourish in the fluid of the inflammatory oedema.
Although the serous exsudation of anthrax does not
usually contain many microbes, it may nevertheless
serve as a nutrient medium for them. Even in animals
refractory to anthrax, the spores of this bacillus, if in-
troduced into the serous fluid of the oedema produced
by a primary inoculation of anthrax, 1 will germinate
and give rise to a new generation of microbes. The
small number of microbes found in the exsudation can-
not therefore be taken as a proof of any bactericidal
action of this fluid.
In human beings tuberculosis is also frequently
accompanied by serous pleurisy in which no bacilli will
be found. But their absence is not due to their having
been killed by the liquid. If they had been present,
they would, as is invariably the case, have attracted the
leucocytes. As the microbe of diphtheria remains
localised at the seat of inoculation, we cannot doubt
that in this disease also there are no bacilli in the
serous exsudation from the very beginning of the
process. The above-mentioned ten cases of Zimmer-
mann were diseases brought about by the streptococcus
pyogenes and the staphylococcus, i.e. by two microbes
which, as Stern 2 has proved, the serous exsudation of
human beings is absolutely unable to destroy. And yet
Zimmermann was unable to find any bacilli in the serous
exsudation which he collected from the above cases.
1 For a description of experiments of this nature on rats, see
Annales de Vlnstitut Pasteur, 1890, p. 201,
2 Zeitschrift f, klin. Meditin, 1890, vol. xviii., p. 62.
LECTURE XI. 175
After this analysis of the facts so far as they are
known, we cannot admit that serous inflammation is a
means used by the organism for killing the pathogenic
organisms. The results obtained with regard to the
bactericidal properties of serous fluids generally are in
confirmation of this conclusion. In spite of all the
endeavours to prove that an active part is played by
these fluids in the destruction of microbes and the
production of immunity, we must recognise the fact
that they are of no importance in this capacity. Further
study of the bactericidal powers possessed by the serum
has shown that these have no connection either with
the phenomena which go on in the organism or with
the production of immunity.
Behring, who was one of the chief originators of the
bactericidal theory of the humours, carried out a large
number of researches on the subject, and concluded by
doubting whether the blood deprived of its cellular
elements could really produce immunity. 1 The results of
numerous experiments which he made with Nissen 2 led
him to consider that the bactericidal property of the
serum is only met with in a few isolated cases, where
natural or acquired immunity exists at the same time.
Among these cases, the most marked are the action of
the serum of rats upon the anthrax bacillus and that of
the serum of vaccinated guinea-pigs upon the Vibrio
Metchnikowii.
Although the facts discovered by Behring and Nissen
are perfectly correct, and we must regard it as esta-
blished that the serum of rats has really a bactericidal
action on anthrax bacilli, and the serum of vaccinated
1 Deutsche medicin. Wochenschrift, 1891, No. 19, p. 655.
2 Zeitschrift fur Hygiene, 1890, vol viii. p. 424.
176 INFLAMMATION.
guinea-pigs on the Vibrio Metchnikowii, yet even in
these cases these bactericidal powers which are so
evident in vitro do not come into play in the body of
the living animal. As Hankin, 1 Roux and 1 2 have
shown, although rats are not refractory to anthrax, yet
their serum outside the body has a bactericidal action
on the bacilli of this disease. Again the Vibrio Metch-
nikowii is readily destroyed in vitro under the influence
of the serum of guinea-pigs which have been rendered
refractory by vaccination, although if inoculated under
the skin or in the eye of the same animals, these
microbes live for some time and finally adapt themselves
to their environment, so that they can now live in the
serum prepared from these animals. 3
The impossibility of drawing any conclusion as to
the events that occur in the living organism, from the
results obtained from experiments with the serum out-
side the body, was shown in the earlier researches of
Lubarsch 4 and myself. 5 Recent experiments of
Buchner, Ibener, and Rceder, 6 have also proved that
the bactericidal properties of the serum, which are
very evident when the serum is allowed to act upon the
microbes in test tubes in the usual manner, become
much less marked if the same microbes be introduced
into the living body enclosed in capsules of filter paper,
so that they may be acted upon by the fluids of the
body without the possible interference of cells.
' Centralblatt fur Bacteriologie, 1891, p. 378.
2 Annales de FInstitut Pasteur, 1891, p. 479.
3 Ibid., p. 465.
4 Centralblatt fur Bacteriologie, 1889.
5 Virchow's Archiv, 1888, vol. cxiv. p. 472.
6 Munchener medicin. Wochenschr., 1891, Nos. 32 and 33.
LECTURE XI.
A review of these facts leads us to conclude that the
exsudation of a serous fluid in inflammation cannot be
regarded as a means which the organism may make use
of to destroy the pathogenic microbes, this service being
performed essentially by the phagocytes. Since, how-
ever, the poisonous chemical products of the bacteria
are the most important agencies in producing the general
intoxication that we know as the disease, it is possible
that the serous exudation may serve to attenuate or
modify the action of these products.
The remarkable discovery of Behring and Kitasato l
that the serum of immunised animals possesses antitoxic
properties, seems to tell in favour of this hypothesis.
These observers who have been confirmed by Vaillard,
Tizzoni, and Cattani, have shown that the serum of
rabbits which have been vaccinated against tetanus, can
destroy considerable quantities of the toxine of this
disease. Behring 2 has obtained analogous, though not so
perfect, results for diphtheria ; and the Klemperers 3 have
also proved that the blood and serum of rabbits which
have been inoculated against pneumonia, as well as these
fluids in men who have survived the crisis of a pneumonic
attack, are endowed with antitoxic properties.
Is it possible that the serous exsudation, although in-
capable of killing the microbes, may serve to destroy
the toxines ? This supposition would seem to receive
some support from the discovery by Behring of the
diphtheritic poison in the pleuritic effusion of rabbits
affected with this disorder. But in man, on the other
1 Deutsche medinische Wochenschrift, 1890, p. 1113.
2 Jbid., p. 1145.
3 Berliner klinische Woclienschrift, 1891, Nos. 34 and 35.
N
178 INFLAMMATION.
hand, although diphtheria often ends favourably, the
production of serous exsudations is never observed.
If we may regard croupous pneumonia as a disease
which terminates by producing an antitoxine (although
this cannot be taken as proved, in spite of the statements
of the Klemperers), it would furnish us with another in-
stance of the destruction of a toxine occurring indepen-
dently of any serous inflammation.
Tetanus is the type of a toxic disease. Yet although
vaccination against it gives rise to a rich production of
antitoxine, we never find any serous exsudation occur-
ring in the disease itself. Vaillard, who has a large
experience in the subject of tetanus, informs me that
in no case are serous inflammations ever present either
in animals affected with tetanus, or in those which are
undergoing the process of immunisation by the injec-
tion of the serum of vaccinated animals.
On the other hand tuberculosis, which is so often
accompanied by serous exsudations, must be looked
upon as one of the diseases in which toxines play the
smallest part. This is shown by the fact, among others,
that the guinea-pig, which is so susceptible to tuber-
culosis, is remarkably insensible to the action of the
tuberculin. As another example I may cite the Vibrio
Metchnikowii. If this microbe be introduced into
guinea-pigs, whether susceptible to or vaccinated
against its action, an abundant serous exsudation is
formed, and yet no antitoxine is produced in the animals
that recover from the disease, as is shown by the fact
that these animals, even when vaccinated against infec-
tion by -this microbe, are still susceptible to the action
of the toxine formed by the Vibrio.
Thus the hypothesis that a serous inflammation is a
LECTURE XI.
salutary reaction of the organism, is not confirmed by
the facts so far as we know them. We must keep in
mind, however, that our knowledge on the subject of
the production of antitoxines is in its infancy ; and we
must wait for much wider and more varied experience
before we can come to a conclusion on the subject.
I may mention one more hypothesis. If we cannot
look upon the exsudation of serous fluid as connected
with the formation of antitoxines, it is at least possible
that it may serve to dilute the bacterial toxines and so
render them less active.
A study of comparative pathology shows that in the
evolution of the inflammatory process the serous in-
flammations are much more recent than those which
are merely attended with a leucocytic reaction. In
none of the invertebrata, in which we have studied the
accumulation of phagocytes around foreign bodies, is
there any trace of a serous exsudation accompanying
this process. This is true not only for the organisms
which possess no blood (such as the Sponges, Ccelente-
rata and Echinoderm larvae) but also for those en-
dowed with a circulatory system. Even in the amphi-
bians, in which the phagocytic reaction is so pronounced,
the inflammation is unaccompanied by any marked
transsudation of fluid. Only in a few exceptional cases
have I observed a slight accumulation of fluid around
inflammatory foci in the tails of tadpoles and of urodelan
larvae.
From whatever point we consider serous inflamma-
tion it appears as a phenomenon of minor importance
compared with the typical inflammatory process, i.e. an
accumulation of phagocytes in the inflamed area.
N 2
LECTURE XII.
Review of other theories of inflammation in light of the acquired
facts Nutritional theory of Virchow Vascular theory of
Cohnheim Experiments of the latter on the tongue of the frog.
Introduction of irritative agents into the blood. Argument
against Cohnheim's theory furnished by the reaction in verte-
brata Struggle of the organism with external agencies Use
of intracellular digestion Phagocytes Hsemitis (recurrent
fever, disease of Daphnia) Tuberculosis. Essential nature of
inflammation Sensibility of the phagocytes Its progressive
development Sensibility of the endothelial cells Definition
of inflammation. Inflammation is not regeneration Inflam-
mation is not identical with resorption Objections raised to
the biological theory of inflammation Vitalism Teleology
Absence of phagocytes in certain infective lesions Imperfec-
tions in the inflammatory reaction Surgical interference
Comparative pathology.
HAVING now reviewed the principal phenomena of
the inflammatory reaction in the animal kingdom, we
may ask how far the facts thus brought out are in
harmony with the various theories mentioned in the
first lecture.
It is unnecessary to insist more than cursorily on the
impossibility of explaining these facts by the nutritional
theory of Virchow. 1 This doctrine assumes that a
greatly increased flow of nutritive substances takes
place towards the inflamed part as well as abnormal
1 " Cellularpathologie," 4th Edition, 1871, p. 475.
LECTURE XII. l8l
proliferation of the local cells, that in fact inflammation
begins from the moment that nutritional derangement
occurs. This derangement consists in " the attraction,
that is to say, in the direct absorption and the modifica-
tion, according to circumstances, of large quantities of
nutrient substances." In this way the cells of the
inflamed organ receive an excessive amount of nourish-
ment at the expense of the fluid parts of the blood.
Considering the matter from this point of view, Virchow
regards the most characteristic phenomena of inflamma-
tion, not as a salutary means of reaction, but as a
process distinguished chiefly by its acute nature and
especially by the danger that the organism incurs from
it. 1
Samuel and Cohnheim held that inflammation con-
sisted essentially in a molecular lesion of the vascular
wall, that the latter, modified by some injurious agency,
lost its power of retaining the blood corpuscles, which
therefore in consequence of the forces acting upon them
were pressed out of the vessels, and then wandered to-
wards the part of least resistance. According to them,
inflammation was in nowise a mode by which the
organism reacted to extrinsic influences but was merely
a primary lesion of the blood-vessels. One experiment
of Cohnheim's well illustrates his way of looking at the
subject. If by means of a ligature the circulation of
the blood in a frog's tongue be stopped for forty-eight
hours, and the ligature be then removed, circulation
will again proceed, but accompanied with a condition
of inflammation, i.e. with the peripheral distribution of
the leucocytes, followed by their diapedesis. Cohnheim
explains this fact as a direct result of the lesion to the
1 Loc. cit., p. 399.
1 82 INFLAMMATION.
vessel-wall, consequent upon its previous prolonged
state of anaemia. " I consider that the cause of
inflammation, "he adds, "should undoubtedly be sought
in the vessels themselves ; the events outside the vessels
appear to be of too subordinate a character to afford an
explanation of this phenomenon." l And yet very
important events must take place outside the vessels.
The peripheral tissues, deprived of their nutrition and
of their protection by the blood, must become the seat
of aggression on the part of the numberless microbes
which exist in the buccal cavity. The tissues them-
selves, or some of them, must at the same time undergo
degeneration, which may produce sufficient peripheral
excitation to bring about an inflammatory reaction.
The instance given by Cohnheim enables us to com-
prehend his theory without however proving it.
In order to produce what we may call a central
vascular lesion independently of the lesion in the other
organs situated at the periphery, we may adopt a simple
expedient, which consists in introducing the irritating
object inside the vessels themselves. In the same
work of Cohnheim on embolism, he adduces experi-
ments which were made with the purpose of producing
embolic abscesses. To this end he injected into the
arteries of frogs a number of substances (such as
globules of mercury, powdered cantharides, particles of
putrid meat) which might have been expected to cause
considerable inflammation, but had no effect at all.
Now these substances if introduced under the skin are
followed by a marked reaction. If therefore inflam-
mation were only due to a change in the vessel-walls,
how is the absence of this change to be explained in
1 " Die embolischen Processe," 1873, p. 51.
LECTURE XII. 183
cases where the irritating agent comes in direct contact
with the walls themselves ?
Many other facts may be quoted which give the same
results as the above experiments of Cohnheim. Patho-
genic organisms which readily provoke inflammation
are frequently found in the blood without causing any
exsudation. In recurrent fever the blood is crowded
with spirilla which act directly upon the vessel-wall by
their spiral movements as well as in all probability
by their toxic products. If Cohnheim's theory were
correct, this would cause such an alteration in the walls
as to bring about the passage of the corpuscles outside
the vessels. Nevertheless no inflammation occurs in the
organs during the course of this fever, although the
whole body of the patient is ' inflamed ' in the highest
degree. Many other microbes, such as the anthrax
bacillus, cause intense inflammation when 'inoculated
subcutaneously, whereas inflammation in the sense used
by Cohnheim does not occur at all, when the same
organisms are in the blood. This is again the case
when the tubercle bacillus is injected into the blood,
while if introduced outside the vessels exsudative
inflammation invariably results.
Besides these facts which the theory of the primary
lesion of the vascular wall does not take into account
the whole comparative pathology of inflammation is
an argument against Cohnheim's contention. The
reactive phenomena of the invertebrata show that
infiltration precedes the vascular events in the evolu-
tional history of the process, and that the leucocytes, led
by their sensibility and by means of their amoeboid move-
ments, themselves proceed towards the injured spot
instead of passively filtering through a vessel-wall.
184 INFLAMMATION.
But even if the theories of Virchow and Cohnheim
cannot be accepted in the light of our present know-
ledge, must we renounce all hope of comprehending
imflammatory phenomena, and remain satisfied with a
mere description of them as most modern pathologists
do?
The study of inflammation from the point of view of
comparative pathology proves first of all that this
phenomenon is essentially reactive in its nature. The
organism, threatened by some injurious agency, pro-
tects itself by the means at its disposal. Since, as we
have seen, even the lowest organisms, instead of
passively submitting to the attacks of morbid agents,
struggle against them, why should not the more highly
developed organisms, such as man and mammals, act in
the same manner ? We must conclude then that the
invaded organism fights against the injurious cause, but
in what way ? As the evolution of inflammation shows,
it is this phenomenon itself which is both the most
general and the most active means of defence among
the members of the animal kingdom.
The essential factor in inflammatory reaction is an
endeavour on the part of the protoplasm to digest the
harmful object. This digestive action, in which the
whole or almost the whole organism of the Protozoa
takes part, is undertaken by the entire plasmodic mass
of the Myxomycetes, while, from the Sponges upwards,
it is confined to the mesoderm. In those cases where
the victory remains with the invaded organism, the
phagocytic cells of this layer assemble, englobe and
destroy the injurious agent. This phagocytic reaction,
in the lower scale of animal life, is slow, owing to the
progression of these cells towards the injurious body
LECTURE XII. 185
being dependent solely on their amoeboid movements ;
but as soon as a circulatory or vascular system makes
its appearance in the course of evolution, it becomes
much more rapid. By means of the blood-current the
organism can at any given moment send along to the
threatened spot a considerable number of leucocytes to
avert the evil. When the circulation is partially
carried on by a lacunar system there is nothing to
intercept the movement of the leucocytes towards the
seat of injury. But when these cells are enclosed
within the vessels, they are obliged to adapt themselves
specially to fulfil their object, which they do by passing
through the vascular wall.
If we accept this conclusion that inflammation in the
higher animals is a salutary reaction of the organism
and that diapedesis and its accompaniments form part
of this reaction, several details of inflammatory pheno-
mena will appear clear to us. For instance the lobed
and polymorphous shape of the nucleus of the pus-
corpuscles has long been remarked. This particular
shape is peculiar to the polynuclear leucocytes, which
represent the vast majority (75 per cent.) of the
total number of white cells. As it was noticed
that a quantity of pus-corpuscles died in the exsu-
dation, this fact became associated with the curious
form of the nucleus ; it was said, and is still maintained,
that the polynuclear leucocytes are cells predestined to
perish and incapable of any considerable activity. On
the contrary these leucocytes are precisely the most
active cells in the organism. The shape of their nucleus
may be more adequately explained as a special adapta-
tion for passing through the vessel-wall. If the process
of diapedesis be watched, the difficulty experienced by
1 86 INFLAMMATION.
the nucleus in getting through will at once be noticed.
Directly this has occurred, the rest of the protoplasm
follows rapidly. It is obvious that a nucleus divided
into several lobes can pass through the wall more easily
than one not so separated. Hence in pus the poly-
nuclear leucocytes are more numerous than the mono-
nuclear leucocytes, and hence the lobed shape of the
nucleus is found only in the leucocytes adapted for
diapedesis and does not occur among the inver-
tebrata (except in a few Cephalopoda).
The facts which demonstrate the untenable position
taken up by Cohnheim may be readily explained by
means of the theory here set forth. If the irritating
agent be outside the vessels, it provokes a typical in-
flammation, accompanied by diapedesis ; if the same
agent be within the vessels no diapedesis takes place
but the leucocytes fight against the microbes in the
blood itself. For instance in recurrent fever, the
spirilla undoubtedly act upon the vascular wall with-
out bringing about diapedesis. But the leucocytes in-
crease in number ; the leucocytosis is followed by a
struggle which is ended by the leucocytes devour-
ing the spirilla. We have here a case of inflammation
unaccompanied by diapedesis ; the conflict between the
phagocytes and the spirilla takes place in the blood
itself. Although no diapedesis occurs, there is in re-
current fever great elevation of temperature as well as
other symptoms which prove it to be an inflammatory
disease. It is apparently a case of inflammation in the
blood itself, a sort of ' hemit'is ' as Piorry considered it
might be many years ago. We find the same condi-
tions in animals whose vascular system and general
body-cavity are in communication. Thus as we have
seen in the disease of Daphnia, caused by the Mono-
LECTURE XII. 187
spora, the leucocytes often collect in large numbers
around the spores of this parasite their assemblage
taking place in the body-cavity.
As another instance we may cite tuberculosis. If
inoculated subcutaneously, the tubercle bacilli produce
inflammation accompanied by considerable diapedesis.
But if the same bacilli be injected direct into the blood,
no diapedesis occurs, but the phagocytes will gather
round the bacilli within the vessels and form intravas-
cular tubercles. It cannot be said that in the first case
(extravascular inoculation) there is inflammation and
that in the second (intravascular inoculation) there is
none, especially as the same tubercles are formed in
both instances. This is another example of an inflam-
mation of the blood itself.
All these cases of intravascular inflammation without
diapedesis, as well as the inflammatory phenomena in
the young larvae of Axololotls and Tritons (where it is
the migratory cells that collect at the seat of injury),
in fact the whole series of reactive phenomena in so
many of the invertebrates, prove clearly that the essential
and primary element in typical inflammation consists in
a reaction of the phagocytes against a harmful agent. If
the latter be in the general body-cavity, which is filled
with blood, the phagocytes will collect here ; if in the
interior of the vessels, as in recurrent fever or in
intravascular tuberculosis, the phagocytes will assemble
in the blood itself; if on the contrary the injurious
agent is outside the body-cavity or outside the vessels,
the phagocytes will emigrate towards the threatened
spot an emigration without diapedesis in the inverte-
brata and young larvae of Urodela, or with diapedesis
in the vertebrata.
Before phagocytic reaction can take place, these cells
1 88 INFLAMMATION.
must be excited positively. Negative sensibility may
also serve as a means of defence in a mobile organism,
such as the plasmodium of the Myxomycetes, which
retires from the offending cause. In the cases where
the latter has penetrated into the organism, negative
sensibility on the part of the phagocytes will leave the
field of battle to the parasite, so that, as frequently
happens, the death of the organism results. Hence as
we rise in the scale, we are met by a progressive evolu-
tion of positive sensibility in the leucocytes. In
Daphnia the observer is struck by the number of
diseases in which phagocytosis is entirely or almost
entirely absent. By the time we reach the amphibia,
positive chemiotaxis is already very marked and, as
Gabritchewsky has shown, it is still more highly
developed in rabbits. And yet among the rodents, as
in the small laboratory animals generally, there occurs
a certain number of rapidly fatal diseases (such as
chicken cholera, hog cholera, vibrionian septicaemia
of birds), in which phagocytosis is often completely
absent. In man and the higher mammals, similar
diseases are much less frequent.
But, in addition to the mobile phagocytes adapted by
their sensibility to move towards the offending object,
there are also fixed phagocytes. A good example of
the latter, which are especially developed in the higher
vertebrates, is furnished by the endothelial cells of the
vessels. Since these cells are contractile and phago-
cytic, it is natural to conclude that they must also be
possessed of sensibility. Thus, if we assume a chemio-
tactic sensibility of the endothelial cells, we may easily
explain the remarkable power of reciprocal attraction
possessed by the protoplasmic processes of developing
LECTURE XII. 189
capillaries, which enables them to meet and form a new
vascular loop. We may apply this explanation to
account for the fact that in many neoplasms, as in
pannus, the vessels penetrate and branch freely in the
affected tissue, whereas in the granulomata, such as
tubercle, leprosy and actinomycosis, blood-vessels are
absent. In the former case there is a positive chemio-
tactic influence attracting the vascular loops, in the
latter a negative chemictaxis or other form of negative
sensibility of the endothelial cells. The co-operation
of these cells in the inflammatory process would be also
directed by their sensibility, at any rate so far as their
active contraction is concerned.
There is one more form of sensibility that we must
mention, namely that of the nervous system, which aids
the phagocytic and vascular mechanisms in their re-
action against deleterious agents.
To sum up : Inflammation generally must be regarded
as a phagocytic reaction on the part of the organism against
irritants. This reaction is carried out by the mobile
phagocytes sometimes alone, sometimes with the aid of
the vascular phagocytes or of the nervous system.
The theory here indicated might be termed the
biological or comparative theory of inflammation, since
it is founded on a comparative study of the patho-
logical phenomena presented by living cells.
It is above all necessary to emphasise the fact that
the essential phenomena of inflammation represent an
actual struggle between the phagocytes and the irritant
agent. As we have seen that leucocytes are capable of
transformation into connective tissue cells, it might be
thought that this was the only object for which the
accumulation took place. This idea however is nega-
INFLAMMATION.
lived by the fact that, in the higher vertebrates, the
polynuclear leucocytes do not contribute to the forma-
tion of granulation-tissue, although they constitute the
large majority of the cells taking part in the inflamma-
tion. The mononuclear leucocytes are the only cells,
besides the endothelial and connective tissue cells, that
are actively engaged in the formation of the new
connective tissue. These leucocytes are formed
directly from the numerous lymphocytes which flock
towards the locality where regeneration is going on.
It has been often thought that the leucocytes which
are gathered together in an inflamed area may only
serve to effect the absorption of dead cells and microbes,
and they have been looked upon as simple ' scavengers '
of the organism. We have already seen that this
hypothesis is not justified by facts, and that from the
very onset of infection the leucocytes wander towards
and englobe the parasites in a living condition. In
case these direct proofs may not be thought sufficient,
I may mention other considerations which tell against
this theory.
If the main object of the leucocytic reaction in in-
flammation were the absorption of solid matters, we
should expect to find very pronounced inflammatory
phenomena in all cases where a very considerable and
rapid absorption is going on. This is however not the
case. The metamorphosis of Batrachians is accom-
panied by an absorption of the larval organs tail and
gills. This is effected very rapidly (in a few days) by
the aid of phagocytes which devour all the tissues no
longer required by the animal. And yet in this case
there are no signs of inflammation, the phagocytes of
the tissues in question being quite sufficient by them-
LECTURE XII. 191
selves to carry out the work of absorption which is evi-
dently a much easier task for them and requires much less
activity on their part than does the struggle with parasites.
In mammals the real ' scavengers,' that is to say the
phagocytes which carry out the work of absorption, are
the macrophages, especially the mononuclear leucocytes.
The latter cells play an important part chiefly in the
chronic inflammations, such as tuberculosis, while in the
acute inflammations it is mainly the microphages, or
neutrophile polynuclear leucocytes, that are involved-
In erysipelas for example the streptococci are englobed
only by the polynuclear leucocytes, and are never
taken up by the macrophages, which, however, carry
out the entire work of absorption, and even cnglobe the
microphages, many of which perish in the struggle with
the microbes and have to be themselves absorbed. If
absorption were the only function of the inflammatory
emigration, it would be teleologically absurd for this
process to end in the formation of pus, i.e. a mass of
leucocytes, many of which die and must themselves be
absorbed. It is much more natural to assume that the
emigration is a reaction on the part of the organism
and that in the conflict many of the principal com-
batants, the microphages, perish on the field of battle.
Absorption afterwards ensues and is carried out by
another variety of phagocytes.
When I first put forward the biological theory of
inflammation eight years ago, 1 I expressed the idea
that this reaction is effected by the intermediation of a
physiological continuity between " the cells of the
connective tissue, those of the endothelial wall and the
leucocytes, which form a complete chain and play the
1 Biologisches Centralblatt, 1883, p. 564.
192 INFLAMMATION.
principal part in the inflammation of vertebrates."
The connective tissue cells which are first attacked,
would, I thought, transmit the action to the vascular
wall, the cells of which would contract to facilitate the
passage of the white corpuscles. At that time only
tactile sensibility of these various cells had been recog-
nised, although certain facts, such as the existence of
chemiotaxis, was already suspected from observations
of the power possessed by the sexual cells of the
Hydromedusae to travel considerable distances. 1 Since
then this chemiotaxis has been definitely proved to exist.
This biological theory has often been considered too
vitalistic in its tendency. I need only quote Frankel's
outspoken criticism of my theory from this point of
view. 2 " The phagocyte theory presupposes extra-
ordinary powers on the part of the protoplasm of
leucocytes, to which are attributed sensations, thoughts
and actions, in fact a kind of psychical activity." The
sensibility of the phagocytes is not an hypothesis which
can be admitted or rejected at will, but an established
fact, which cannot be ignored, as it is by Frankel.
Whether they possess powers of thought and volition,
as this author accuses me of assuming, is quite beside
the question, though we are justified in considering that
they possess a germ of these qualities and that their
sensibility, like that of various vegetable and animal
unicellular organisms represents the lowest stage in the
long series of phenomena which culminate in the
psychical activities of man.
1 Weismann, " Die Enstehung der Sexualzellen bei Hydromedu-
sen." Jena, 1883.
1 "Grundriss der Bakterienkunde." 3rd Ed., Berlin, 1890,
p. 203.
LECTURE XII. 193
As the authorities on these subjects, Herbert Spencer,
Romanes and others have shown, psychical phenomena
have nothing specific in them, but have developed as an
association of the simple actions which we observe in
the lower organisms and the cells of different animals.
The accusation of vitalism and animism, which is
unjustly cast at the phagocyte theory, might really be
more appropriately applied to my opponents, who
maintain that the psychical acts of the higher animals are
fundamentally different in their nature from the more
simple phenomena peculiar to the lower organisms.
It is equally erroneous to attribute a teleological
character to the theory that inflammation is a reaction
of the organism against injurious agencies. This theory
is based on the law of evolution according to which
the properties that are useful to the organism survive
while those which are harmful are eliminated by natural
selection. Those of the lower animals which were
possessed of mobile cells to englobe and destroy the
enemy, survived, whereas others whose phagocytes did
not exercise their function were necessarily destined to
perish. In consequence of this natural selection the
useful characteristics, including those required for
inflammatory reaction, have been established and trans-
mitted, and we need not invoke the assistance of a
designed adaptation to a predestined end, as we should
from the teleological point of view.
But, it has been urged on several occasions, 1 if the
phagocytic reaction has been developed in order to
protect the organism from danger, how is it that the
phagocytes refuse to act just when the organism is most
1 For instance by Baumgarten, Berlin, klin. Wochenschrift, 1884,
and Burden Sanderson, Brit. Med. Journ., 1891, p. 1085.
O
1 94 INFLAMMATION.
threatened ? This objection again arises from an
insufficient knowledge of the principles of the theory.
It is just because the defence by the phagocytes is
developed according to the law of natural selection and
is not a designed adaptation to a particular end, that
cases naturally occur where the phagocytes do not
fulfil their functions, a neglect followed by the most
serious danger to or death of the organism. In nature
the organism is possessed of many characteristics, which
may be either useful or injurious to their owner. The
former causes the survival, the latter the death of the
possessor. Let us take two organisms : one in which
the phagocytes are readily repulsed by the microbe, the
other whose phagocytes show a positive sensibility
causing considerable phagocytosis. The former will
soon fall a prey to the parasite and be eliminated by
natural selection, whereas the latter will resist the
infection, survive and put forth progeny possessed of
the same phagocytic properties. Under these con-
ditions the activity of the phagocytes will increase
with every successive generation.
But the curative force of nature, the most important
element of which is the inflammatory reaction, is not
yet perfectly adapted to its object. The frequency of
disease and the instances of premature death are a
sufficient proof of this. The phagocytic mechanism
has not yet reached its highest stage of development
and is still undergoing improvement. In too many
cases the phagocytes flee before the enemy or destroy
the cells of the body to which they belong (as in the
scleroses). It is this imperfection in the curative forces
of nature which has necessitated the active intervention
of man.
LECTURE XII. 195
The defence of the organism against deleterious
agencies, which is at first confined to the phagocytic
mechanisms and the somatic system of nerves, by and
by spreads to and is undertaken by the psychical
nervous apparatus. With the nervous cells which
direct the contraction and dilatation of the vessels
become associated other cells which control thought
and voluntary actions. One function of these psychical
cells has been to develop a complete science for the
defence of the organism against hostile influences. By
its means, methods for assisting the curative inflamma-
tion have been devised, as for instance the formation of
artificial lesions in order to facilitate the inflammatory
reaction. The application of agents which set up in-
flammation, such as jequirity, the virus of gonorrhea,
tuberculin and cantharidin, is the conscious continua-
tion of the defensive measures which have been uncon-
sciously evolved by the long series of animals in their
struggle for existence.
But neither the conscious nor the unconscious
mechanism is perfect. Medical science should therefore
call to its aid all the less complex branches of science,
including that of biology, which seeks to investigate
organisms and their evolution. Not only inflamma-
tion but many other medical problems might be advan-
tageously attacked from the standpoint of comparative
pathology. In order to trace the first appearance of
febrile reaction, the study of fever should begin with in-
vestigations on heat production in the lower animals, con-
tinuing on to the vertebrates, such as reptiles, &c. As
several of these phenomena have originated at an early
geological epoch, the conditions in which the first warm-
blooded animals lived should also be taken into account.
O 2
196
INFLAMMATION.
Comparative pathology could undoubtedly throw
light on the important pathological question of tumours-
As many of the lower organisms both animal and
vegetable are subject to the formation of tumours, the
part played by parasites in their etiology could be
more easily established and the theory of embryonic
aberrations refuted.
The subject of atrophy, intimately connected with
that of chronic inflammation, is likewise a suitable one
for the application of comparative pathology to its
elucidation. Here again the phagocytic phenomena
take a prominent part
If however medical science may learn much from
biology, of which it forms but a part, it may at the
same time give something in return. General biology
may extend its knowledge by including the study of
the morbid phenomena of which pathology takes cog-
nisance. In biology difficulties frequently present them-
selves in the study of the evolutionary processes, be-
cause the perfect condition of adaptation in which these
phenomena occur in nature, is not a favourable one for
analysis. In order to throw light upon the universal
law of natural selection, it is necessary to study the
less stable phenomena, the less perfect mechanisms, in
a word, the phenomena in which the working out of
this law can be watched day by day. Now it is just
the morbid phenomena with the reactions they provoke,
the struggle between the organism and its aggressors,
which offer the best opportunity for a consecutive study
of the course of natural selection. In this struggle
every day some of the combatants, by virtue of natural
selection, survive, while others, eliminated by the same
means, perish. Now it is the conquering organisms
LECTURE XII. IQ7
which survive and the vanquished parasites which be-
come eliminated, now the defeated organisms which are
eliminated and the triumphant parasites which sur-
vive.
Thus I conclude as I began. General pathology
should go hand in hand with zoology or rather with
biology, and form one branch of it, that of comparative,
pathology. This science is only in its infancy, and yet
it is already in a position to render good service to
medicine. By facilitating the analysis of the reactive
phenomena, it indicates the elements which should be
especially protected and reinforced in the conflict of
the organism against its enemies, and thus contributes
to the solution of one of the great problems of
humanity.
DESCRIPTION OF THE PLATES.
E PLATE I.
FlG. I. Inflamed spot on the caudal fin of a larva of Bombinator
igneus, 72 hours after cauterisation with nitrate of silver : a. Pha-
gocytes containing pigment and red corpuscles ; b, c. Branched
cells enclosing fragments of red corpuscles ; d. A phagocyte inside
a lymphatic vessel.
FlG. 2. A cyst of the Gregarina of Lumbncus surrounded by a
thick and deformed cuticle, and by a follicle formed of connective
tissue.
FlG. 3. A cyst of the same parasite, with only the remains of
its cuticle left.
FIG. 4. A Rhabditis in the midst of a mass of phagocytes.
c. Thick and deformed cuticle of the parasite.
FlG. 5. Another Rhabditis, enclosed in the mass of phagocytes
of Lumbricus. c. Cuticle of the parasite, consisting of several
concentric layers.
PLATE II.
FlG. I. A cyst of the Gregarina of Lumbricus with its contents
divided into cells. The cyst is surrounded by a follicle composed
of connective tissue.
FIG. 2. A mass of phagocytes of Lumbricus surrounding a
cyst of Gregarina which they have destroyed.
2OO DESCRIPTION OF THE PLATES.
FIG. 4. Example of positive chemiotaxis. A plasmodium of
Didymium farinaceum which has dipped its processes into an
infusion of leaves.
FIG. 3. Example of negative chemiotaxis. The plasmodium of
Fig. 4 receding from a o'l per cent, solution of quinine.
FIG. 6. Another plasmodium of Didymium repulsed by the
hydrochlorate of quinine (o i per cent.)
FIG. 5. The same plasmodium 5 hours after the solution of
quinine has been replaced by an infusion of leaves. The negative
chemiotaxis of Fig. 6 is converted into positive chemiotaxis.
FIG. 7. A leucocyte of the salamander, a. Attraction sphere
(after Flemming).
FIG. 8. A clasmatocyte from the mesentery of Triton tceniatus.
PLATE III.
FIG. i. A cyst of Gregarina surrounded by the phagocytes of
Lumbricus. (Magnification : objective D and ocular 4 of Zeiss.)
FIG. 2. Eosinophile leucocyte from man (after Gabritchewsky).
FIG. 3. One of Ehrlich's cells from a white rat.
FIG. 4. A giant cell from the spleen of Meriones, a. En-
velope of the bacillus ; b. Bacillus of Koch. (The spleen treated
with Flemming's fluid and stained by Gram's method and cosine.
Magnification : objective 1/18, and ocular 3 of Zeiss.)
FIG. 5. A giant cell from the spleen of Meriones, containing
a calcareous body with a double bacillus. (Stained with haema-
toxylin and Ziehl's fuchsin. Same magnification.)
FIG. 6. Another giant cell within which the bacillus may be
seen surrounded with concentric layers. (Treated and stained as
in Fig. 5. Objective 1/18, ocular 2.)
FIG. 7. A giant cell with a calcareous body containing only a
trace of the bacillus b, (Fuchsin, haematoxylin, 1/18x2.)
FlG. 8. Another giant cell, in which the bacillus b has become
converted into a pale rose-coloured body.
FIG. 9. A giant cell enclosing a fully developed calcareous
body.
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APPENDIX.
A CONSIDERATION OF SOME CRITICISMS J ON THE
BIOLOGICAL THEORY OF INFLAMMATION.
IN the foregoing pages the ontogenetic study of the pheno-
mena of inflammation in the animal kingdom has enabled me to
establish a biological theory of this process. I have shown
that it really represents an adaptation of the organism to its
environment, which has been developed in the course of its
never-ceasing struggles against deleterious agencies. " In-
flammation generally must be regarded as a phagocytic reaction
on the part of the organism against irritants. This reaction is
carried out by the mobile phagocytes, sometimes alone, some-
times with the aid of the vascular phagocytes or of the nervous
system." (P. 189.)
In analysing the characteristics of inflammation, pathologists
have long recognised that the exsudation forms the principal
feature, and that the three other classical symptoms, heat,
redness and pain, are only of secondary importance. Now the
most important constituents of the exsudation are the leucocytes,
which are contained in large quantities, not only in purulent
but also in serous and fibrinous exsudations. It is rare to find
them completely absent in any inflammatory exsudation. The
fact, that in the lower animals the inflammatory reaction consists
merely in an aggregation of phagocytic cells similar to leuco-
cytes without any accumulation of the body fluids, shows that
the existence in higher animals of exsudations free from cells
is of secondary importance for our conception of the essential
1 Podwyssozki. " Memoir in Commemoration of Virchow's
Jubilee," Kieff, 1892 (in Russian). Ziegler, " Historisches und
Kritisches iiber die Lehre von der Entzundiing," b'ettrage zur
pathologischen Anatomte, 1892, vol. xii., p. 152, Weigert, Deutsche
medicinische Wochenschtift, 1893, pp. 17, 37.
202 APPENDIX.
nature of the process. It is evident however that these
phenomena must be taken into account in formulating any
scientific theory of inflammation.
The criticism usually urged against the biological theory of
inflammation has been that it totally disregards inflammations
in which the exsudation is free from leucocytes. According to
Podwyssozki, certain serous inflammations may occur un-
accompanied by any phagocytic reaction, and yet we cannot
exclude these from the category of true inflammations. He
proposes therefore to replace the above definition by the
following. " Inflammation is a local reaction, often beneficial,
of the living tissues against the irritant substance. This re-
action is chiefly produced by a phagocytic activity of the meso-
dermic cells. In this reaction however may participate not
only changes in the vascular system, but also the chemical
action of the blood-plasma and tissue-fluids in liquefying and
dissolving the irritant agent." This theory has the advantage
of including the purely serous inflammations.
In the same way Weigert accuses me of ignoring the
existence of exsudations free from cells, and suggests that I do
not know what to make of the serous inflammations. Ziegler
raises the same objection, although he does not formulate it so
precisely. These criticisms however rest upon a misconcep-
tion which we may remove at once. Although I have devoted
a volume to the development of the biological theory of in-
flammation, I have had to leave many points insufficiently
discussed, and it is just to these points that the principal
objections have been urged.
The passage of red corpuscles and plasma into the inflamma-
tory exsudations must be looked upon as the result of the
activity of the endothelial cells of the vessel walls. (P. 149.)
The frequent presence of red corpuscles in the exsudations
which are most serous and free from leucocytes shows clearly
that in these cases there is a direct passage of the constituents
of the blood into the exsudation. If the blood discs are able
to pass through the vascular wall, it is evident that the blood
plasma must undergo the same fate. It is impossible therefore
APPENDIX. 2O3
to ascribe the serous exsudation to a secretory activity on the
part of the endothelial cells. These cells on the contrary
must permit of the passage of the plasma and corpuscles in
consequence of the contractility which is one of their pro-
perties. We may compare this phenomenon to those observed
in the Sponges, where the ectodermic cells by their contraction
open or close the apertures of the canals and so permit or
refuse the passage of fluids with the bodies they may hold in
suspension. In both cases, the passage of water through the
ectodermic wall in the Sponge, and of plasma through the
endothelial wall in inflammation, are brought about in con-
sequence of the sensibility and contractility of the cells
concerned. Although the process cannot be called phagocytic
in the strict sense of the term, yet its intimate relationship with
phagocytic phenomena is very evident.
Phagocytosis is a phenomenon of considerable complexity.
When it is exhibited by leucocytes, these cells are in the first
place affected by various substances which possess an attrac-
tion for them. They proceed towardsthese substances by means
of their amoeboid movements and then englobe them. Intra-
cellular digestion may afterwards occur. Here then we have
phenomena of sensibility, contraction, ingestion, and produc-
tion of digestive fluids. As a matter of fact the sequence is
frequently broken at some point. Thus when a guinea-pig is
infected with anthrax, the leucocytes, acted upon by the
bacterial products, approach the microbes ; leucocytosis takes
place, but phagocytosis is arrested, and the bacteria are either
not englobed at all, or to a very slight extent. In this case
there is a phagocytic reaction, which however falls short of
the mark.
In the most complete phagocytic reaction, all the varieties
of phagocytes englobe and destroy the irritant bodies. In
other instances it is only the mobile phagocytes which fulfil this
function. In a third set of examples, the phagocytic reaction is
still less complete. The leucocytes remain in the organs and
in the blood and do not pass over into the exsudation ; the en-
dothelial cells alone react, but do not accomplish all the phases
204 APPENDIX.
of phagocytosis ; they stop short at a stage of contraction which
allows the plasma and the red blood corpuscles to pass through
the vessel wall. The most familiar examples of this incomplete
phagocytic reaction are met with in certain very acute diseases
produced experimentally. It is in these cases that purely serous
inflammation has been most studied. Thus in the acute diseases
produced in guinea-pigs by injection of vibrio Metchnikowii or
in rabbits by the coccobacillus of hog cholera when death ensues
after a few hours, the phagocytic reaction is limited to this
condition of contraction of the vascular endothelial cells, giving
rise to a serous exsudation in the infected regions. Where
the disease takes a still more rapid course, as in the most
malignant forms of chicken cholera in the rabbit, no exsuda-
tion at all occurs. There is no phagocytic reaction whatever,
but at the same time there is no inflammation.
We see then that serous inflammation is included perfectly
well by the general conception of the biological theory without
modification. In formulating this theory, I have, in order to
make it as short as possible, only mentioned ' phagocytic re-
action ' in general terms. I have not alluded either to the
sensibility or to the contractility of the phagocytes, since these
phenomena are already comprised in the one notion of the
phagocytic reaction. For the same reason I have not especially
referred to the sensibility and contractility of the endothelial cells,
which are certainly phagocytic in their nature, although these
phenomena play an important part in the production of the
serous exsudation.
Since the comparative pathology of inflammation has
established the reactive and beneficial character of this pheno-
menon, we should expect to find certain close affinities between
it and other processes of the organism. Thus inflammation is
connected by a whole series of intermediate conditions with
other phagocytic phenomena, such as the passage of leucocytes
through mucous membranes ; and chronic inflammation is inti-
mately associated with the atrophy of certain tissues. In fact,
from whatever side we may consider inflammation, we shall
always find connecting links with other natural phenomena.
APPENDIX. 205
Thus from the purely clinical standpoint, no hard and fast
boundary line can be drawn between inflammation and hyper-
asmia, all the intermediate stages between these two conditions
being met with at some time or another.
II.
The severest critic of the biological theory of inflammation
is Professor Ziegler, who considers that it is quite erroneous to
attribute a fundamental importance to phagocytosis in inflam-
mation. I regret that limits of space prevent my reproducing
here all his arguments, and must content myself with quoting
his principal objections. He writes, " Metchnikoff affirms
quite arbitrarily that the pathological phenomenon which
interests him represents the essential part of inflammation. He
is moreover inconsequent in his statements, for he sometimes
regards the phagocytosis exercised by leucocytes and sometimes
the accumulation of mesodermic cells as the essential feature."
He continues, " I look upon the phagocytosis which occurs in
the course of an inflammation as a purely accidental pheno-
menon, which is often brought about for the simple reason
that mobile cells happen to be present, together with a material
capable of being ingested by them." (Ziegler, loc. tit., p. 200.)
When we analyse inflammation by means of a study of its
ontogeny, we are inevitably forced to the conclusion that
phagocytosis is the most primitive phenomenon of the reaction
against irritant agents. Inflammation in cold-blooded animals
teaches us that increased temperature is not a necessary
factor; and the analogous reaction in invertebrates proves
that inflammation may occur without any intervention on
the part of the blood-vessels. The phenomena thus become
more and more simple as we descend the animal scale, till
finally the sole phenomenon we have to deal with is phagocy-
tosis. Since the leucocytes are derived from the mesoderm,
there is no inconsequence in admitting the existence of a
leucocytic phagocytosis and a mesodermal phagocytosis. The
facts are so clear that there would be no advantage in dwelling
on the matter at any greater length.
206 APPENDIX.
Ziegler continues, " When any bodies, such for instance as
bacteria giving rise to attracting substances, are present at any
spot in the tissues, the leucocytes move towards these bodies,
and in some cases manage to englobe them. When on the
contrary the foreign bodies, such as bacteria, exercise a repel-
lent or paralysing effect, the cells will move in an opposite direc-
tion, or remain where they are. The emigration and pha-
gocytosis then are not brought about by the combative
ardour of the cells, but by the properties of the foreign body
which has been introduced, or of the tissues and tissue-juices
modified by the irritant." He concludes, " The idea that
the characteristic feature of inflammation is a struggle of
phagocytes must therefore be rejected." (loc. fit., p. 202.)
Ziegler forgets that the attraction and repulsion of leucocytes
depend not only on the products of the microbes but also on
the sensibility of the leucocytes. Substances which repel the
leucocytes of susceptible animals attract on the contrary those
of animals which are immune by nature or rendered refractory
by vaccination. This fact is so general and well established
that no one is justified in ignoring it. It is precisely because
on the one side we have microbes which defend themselves or
attack by means of their toxic products, and on the other
phagocytes which approach and englobe the microbes, that I
have formulated the idea of a struggle between two living
organisms. None of Ziegler's objections can overthrow this
interpretation of the facts just mentioned.
Ziegler proceeds to make use of an argument which has
often been brought forward. "In certain cases the phago-
cytosis exercised by the leucocytes may help to destroy the
foreign bodies. In other cases however the phagocytosis may
aid in the generalisation of an infectious disease, e.g. when
the bacteria can multiply within the cells, as in leprosy, or
when they are carried about by the cells." (P. 202.) It cannot
be denied that the phagocytic reaction is far from representing
a perfect mechanism, as is evidenced by the frequency of many
diseases. But there is also no doubt that the generalisation
of bacteria takes place much more rapidly in the cases when
APPENDIX. 207
they are not englobed by the phagocytes. Thus one often
hears that a seat of injury may become infected by tubercle
bacilli which have been carried there by leucocytes; but
it must not be forgotten that in tuberculosis the leucocytes
serve to localise the disease and prevent the dissemination of
the bacilli. The importance of the leucocytes as carriers of
infective bacilli has been very much exaggerated.
Not approving either of the biological theory of inflammation
or of the comparative method on which it is based, Ziegler
defines " inflammation as a local tissue-degeneration associated
with pathological exsudations from the blood-vessels." (loc.
cit. t p. 173.) This definition is nothing but a statement
of a certain number of the phenomena of inflammation, and
does not touch the essential factor in this process. The
accumulation of migratory cells in Urodela round irritant
bodies, which, as I have shown, takes place without any
affection of the vessels, as well as all the analogous phenomena
in invertebrata, are quite excluded by Ziegler's definition.
And yet any scientific definition should take account of the
natural affinities which undoubtedly exist between these
phenomena and the inflammation accompanied by a vascular
reaction. Another still more striking illustration of the
insufficiency of this definition is afforded by tubercle. A
tubercle formed within the vessels would not come under
Ziegler's definition of inflammation, whereas a tubercle which
has developed outside the vessels would be a true inflamma-
tion. Now it cannot be denied that intravascular and extra-
vascular tubercle represent essentially one and the same
pathological formation. I must conclude then that Ziegler's
definition, which leaves the main question untouched and
takes no account of natural affinities, must be rejected.
III.
With the exception of the question of serous inflammation,
discussed in Section I., Weigert merely makes a few critical
remarks with regard to certain subsidiary points in the
biological theory of inflammation. My learned critic expresses
208 APPENDIX.
doubts as to the justice of comparing the destruction of the
microbes within the phagocytes to intracellular digestion. He
thinks that there is a contradiction in the fact of intracellular
digestion taking place in an acid medium in Protozoa while
the processes in the phagocytes are carried out in a neutral or
alkaline medium. But besides the case in which the phago-
cytes show an acid reaction (exsudation-cells of the tail of
tadpoles), I may instance the intracellular digestion in multi-
cellular animals. In Actinia this takes place in an acid medium,
in the Spongillse in a neutral or alkaline medium. And yet these
two examples are closely allied to each other. It should be pre-
mised as a general rule that intracellular digestion may present
great variations, and may proceed in media of varying reaction.
Weigert finds another contradiction between my opinion that
bacteria are destroyed within the phagocytes and my men-
tion of the ferment of leucocytes, discovered by Leber,
which acts outside these cells (a ferment which peptonises
gelatin). Here a misunderstanding has arisen. I have never
asserted that the phagocytic destruction of bacteria is carried
out by means of any ferments, still less by means of those
which peptonise gelatin. I have always openly acknowledged
that the question as to what substances within the phagocytes
harm and destroy the microbes is still quite undecided.
They may be ferments, digestive or otherwise, or they may be
substances, acid or alkaline, completely different from fer-
ments. We shall have to find new and more perfect methods
before being able to solve this delicate problem.
In the question of giant cells, Weigert remains faithful to
his old theory, which has been discussed in the " Annales de
1'Institut Pasteur," 1888, p. 604. Unfortunately he has not
chosen to discuss this subject in his criticism. This is the
more to be regretted, since the discovery of the special resis-
tance offered by the giant cells of the Algerian rat to the
tubercle bacillus might give rise to an interesting change of
opinion. It is unnecessary to insist in greater detail on the
fact that this discovery is an important confirmation of my
views as to the giant cells being a means of phagocytic defence.
APPENDIX. 209
IV.
Having 'now replied to the principal objections made to
the biological theory of inflammation, there is one point of a
different nature left to deal with. In spite of his opposition to
the phagocytic theory, Ziegler lays a certain claim to priority
in this question. I may take this opportunity of replying to
this attack as well.
To quote Ziegler's own words : " the phenomenon of phagocy-
tosis has been known for a long time ; in the sixth decade of
our century, experimental researches on the enclosure of coal-
dust and of coloured granules by leucocytes were frequently
made, as well as on transport by these cells. In 1874 I ob-
served that, in granulation-tissue, cells having the characteristics
of leucocytes with divided nucleus, together with red corpuscles,
were englobed and destroyed by large cells. As the result of
my researches I concluded that the material englobed was
assimilated and that an act of nutrition consequently took
place" (p. 197). Ziegler insists upon the fact that "his
researches on intracellular digestion in the mesodermic cells
were published eight years before the works of Metchnikoff,
and that at the time Metchnikoff's first papers on phagocytosis
appeared, this phenomenon was well known to pathologists "
(p.i 99 ).
Ziegler is astonished that I gave no historical references in
my earlier works, and especially in my treatise on inflamma-
tion. But in my first paper * where mention was made of
phagocytes, I quoted " the valuable results of histological and
pathological research on the subject of the phenomena of
absorption in vertebrates." I referred the reader to
Ziegler's text-book of pathological anatomy itself. I did not
dwell particularly on Ziegler's work 2 with regard to the inges-
tion of leucocytes by granulation-cells, for the reason that
1 Arbeiten des zool. Inst. zu Wien, 1883, vol. v. p. 157.
8 " Experimentelle Untersuch. iiber die Herkunft der Tuberkel-
elemente," 1875, p. 68.
210 APPENDIX.
this discovery was made four years previously by Bizzozero, 1
who suggested that the leucocytes found in pus in the interior of
large cells had been devoured by the latter. Later on Ziegler
confirmed this conclusion, but in seeking for the analogies of
this process, compared it to the conjugation of cells, such as
results in the formation of zygospores in Spirogyra or in that
of plasmodia by the fusion of cells, &c. He thus regards the
act of ingestion by the leucocytes as a fusion of cells rather
than as a preliminary to intracellular digestion.
I need scarcely say that in none of my publications have I
laid claim to the discovery of the ingestion of solid bodies by
mesodermic cells, nor have I ignored the large number of
researches which have been made on this subject The
phagocyte theory however is still the theme of so much
debate that I have not had the opportunity of entering fully
into historical details. When it is once firmly established, it
will be time enough to determine the exact part taken in its
foundation by workers such as Panum, Gaule, Roser, &c., who
have much more right than Ziegler to be considered the
pioneers of this theory.
Ziegler is mistaken in thinking that the ingestion of solid
bodies by mesodermic cells necessarily involves the conclusion
that the animal organism possesses, in its phagocytic cells,
(mesodermic and others) a very important means of defence
against pathogenic microbes. In order to establish the fact that
the phagocytes constitute an important defensive mechanism,
it was necessary to prove that the leucocytes englobe microbes
in a living and virulent condition and then injure or destroy
them in some way or other. In order to show the impor-
tance of the phagocytes, it was necessary to prove that their
intervention is of usual occurrence. In this task, I have been
obliged to demonstrate the inaccuracy of the work directed
against the phagocytic theory and carried out in Ziegler's
laboratory. Two of the pupils of this author, Palm 2 and
1 Gaz. med. lombarda, 1871, and Wien. med. Jahresber., 1872,
p. 160.
2 Beitr. z. path. Anatomte, vol. ii. p. 480.
APPENDIX. 211
Rogowitch, 1 have made some researches on malignant pus-
tule in man and ' charbon symptomatique.' Palm came to
the conclusion that in anthrax of man " the cells do not play
the slightest part in the sense of the phagocytosis of Metchni-
koff." Rogowitch expressed the same opinion as regards
' charbon symptomatique ' in several species of animals. And
yet it has been definitely shown that both these attacks were
based on unreliable data. It is at the present time indubitably
proved that the bacilli are englobed in large quantities by
the phagocytes both in malignant pustule in man 2 and in
' charbon symptomatique.' 3 The attack emanating from
Ziegler's laboratory is thus unjustified by facts.
In this reply to the criticisms on the biological theory of
inflammation, I have only considered the more important
objections. None of them seem to me to touch either the
foundation of the theory or the method upon which it has been
built up ; hence I do not think that it can be regarded as having
been in any way injured by the opposition raised to it.
E. METCHNIKOFF.
1 Beitr. z. path. Anafomz'e,vol. iv. p. 291.
2 Karg, Fortschr. d. Med. vol. vi. p. 529, and Lubarsch, " Unters.
iib. d. Immunitat," 1891, pp. 1 1 i-i 14. The latter author summarises
his chapter in the following words : " The existence of phagocytosis
in human anthrax, as well as its parallelism with the course of the
disease and the destruction of the bacilli should be considered as
established beyond doubt by the facts I have brought forward."
It is inconceivable how Roger (Traite" de Medecine de Charcot et
Bouchard, vol. i. p. 555) could interpret Lubarsch's article in a
directly opposite manner and assert that in the cases quoted by
him " there was no connection whatever between the intensity of
the phagocytosis and the evolution of the disease."
3 Annales de Flnstitut Pasteur, 1889, p. 194. Ruffer, Brit.
Med.Journ., May 24, 1890.
P 2
INDEX.
ABSORPTION by leucocytes, 190.
Acinetse, parasitism of, 2, 22.
Actinophrys, 15.
Amoeba, merotomy of, 14.
Amoeboid cells in Sponges, 45.
Amphioxus, absence of reactive phe-
nomena in, 94.
Amphophile leucocytes, 113.
Anguillulides, 73.
Anisoplia austriaca, 89.
Annelida, phagocytosis in, 66.
formation of plasmodia in, 67.
Antitoxic function of serous exsudation,
177.
Antitoxines, 178.
Arachnidse, 88.
Argulus, 78.
Arnold, on stomata, 138.
division of leucocytes, 129.
Arthrobothrys oligospera, 73.
Arthropoda, 75 93.
Ascaris mystax, 73.
Ascidians, diapedesis in, 79.
Astropecten pentacanthus, 60.
Atrophy, 171.
Attraction-sphere, 113.
Aurelia aurita, 59.
Axolotl embryos, inflammation in, 95.
BACILLI, alteration of, in amoebte, 19.
englobed in living condition, 115.
tubercle, degeneration of, 161.
Bacillus pyocyaneus, 121.
Bacteria, ingestion by phagocytes of
Bipinnaria, 64.
Bacteria, luminous, 82.
Bacterial infection of insects, 89.
Bacterium termo, chemiotaxis in, 34.
Balbiani, on bacteria in insects, 88.
on merotomy of Infusoria, 16.
epidemic in Paramsecia, 20.
on " pebrine," 92.
Bardach, 161.
Bary, de, on fungi in caterpillars, 89.
on Peziza scltrotiorum, 41.
Basophile cells, 143.
Baumgarten, criticism of biological
theory, 193.
formation of tubercle, 158.
Bipinnaria asterigera, 62.
Behring, on bactericidal action of
serum, 175.
and Kitasato, on antitoxic proper-
ties of serum, 177.
Billet, 82.
Blood-corpuscles of invertebrata, 76.
white, origin of pus-cells from, 9.
Blood-vessels, development of, 137.
in inflammation, 182.
behaviour in earthworm, 70.
Boinet and Borel, 169.
Bellinger, 121.
Bombinator igneus, 103.
Bordet, 118, 145.
Brault, 170.
views on inflammation, 9.
Bruno Hofer, digestion in amoebse, 125.
Buchner, 156.
on bactericidal properties of serum,
176.
on chemiotaxis, 119.
on inflammation, II.
Burdon Sanderson, criticism of bio-
logical theory, 193.
Biitschli, 73.
epidemic in Paramsecia, 20.
on Acinetse, 23.
CALCAREOUS degeneration of tubercle,
162.
Canu, 81.
Capillaries, contractility of, 138.
Cattani, 177.
Cell-division in inflammation, 9, 10.
214
INDEX.
Ceraochalina gibbosa, 52.
Chaetorceros, parasitic on Bipinnaria,62.
Chambers, ciliated, of Sponges, 46.
Charrin, 153.
Chemiotaxis, 33, 117.
in developing blood-vessels, 189.
negative, 146.
Chklarewsky, 147.
Chloroform, action on leucocytes, 148.
Chondrioderma, plasmodia of, 37.
Chondrosia reniformis, 49.
Chronic inflammations, 157-171.
Chun, fragmentation of nucleus, 167.
Chytridiaceas, parasitism of, 24.
parasitic in Nematoda, 73-
Ciliated chambers, 46.
Cirrhosis, 170.
Clasmatocytes, 143.
Cleonus punctiventris, 90.
Cockchafer, 78.
Ccelenterata, 56 65.
Cohnheim, dilatation of vessels in in-
flammation, 149.
views on inflammation, 6, iSl.
experiments on cornea, 8.
origin of granulation-cells, 131.
Coleoptera, phagocytosis in larval, 78.
Commensalism in Sponges, 53.
Comparative pathology, importance of,
4, 12, 195.
Conidia, parasitic in Daphnia, 83.
Connective tissue, formation of, 132.
its changes in inflammation, 142.
Connective tissue cells, behaviour dur-
ing inflammation, 97.
karyokinesis in, 97.
formation from migratory cells, 1 06.
in tubercle, 158.
Copepoda, 75.
Cordictps militaris, 89.
Cornea, Cohnheim's experiments on
the, 8.
Cornil, on inflammation, 8.
Crustacea, phagocytosis in, 78.
Cuticle, protective function of, 87.
Cyclostomata, leucocytes of, 1 14.
Cyst, formation by Gregarina, 69.
DANILEWSKY, 152.
Daphnia, fungoid disease of, 82.
Delacaze-Duthiers, 78.
Diapedesis, 144, 185.
in Ascidians, 79.
in vaccinated animals, 1 54.
Didymium farinaceum, 35.
Digestion, intracellular, in Sponges, 49.
Diphtheria, 177.
Diptera, 88.
Diseases, infectious, of Infusoria,
1728.
in Amoeba, 17.
in ciliated Infusoria, 20.
Disselhorst, action of quinine, 145.
Doris, 77.
Dust-cells, 1 60.
EARTH-WORM, diseases of, 68.
Ebstein and Nicolaier, 169.
Echinodermata, 56 65.
development of, 60.
Ehrlich, 112.
leucocytosis in phthisis, 168.
Emigration of leucocytes, 144.
Endothelium, development of, 137.
peritoneal, phagocytic nature of,66.
Eosinophile leucocytes, 1 12.
Epidemic in Amoebae, 17.
in ciliated Infusoria, 20.
Epithelioid cells, origin of, 132, 159.
Erysipelas, behaviour of leucocytes in,
117.
leucocytosis in, 122.
Englena, infection of, 24.
Euphorbia Cyparissias, 41.
Evolution of inflammation, 109.
FERMENTS in leucocytes, 123.
Fibroblasts, origin of, 132.
Fishes, 95.
' Flacherie,' 89.
Flagellated Monads, digestion in, 19.
infection of, 25.
Flemming, on attraction-sphere in
leucocytes, 113.
mitotic division of leucocytes, 130.
transformation of leucocytes, 135.
Fluss, 150.
Fraenkel, criticism of biological theory,
192.
Frank, lesions in plants, 39.
Fungi, parasitic in Nematoda, 73.
parasitic on Talitrus, 81.
GABRITCHEVSKY, sensibility of leuco-
cytes, 119.
Gamaleiia, 154.
Gasteropoda, spleen of, 77.
Gasteropteron, 77.
Gastrasa theory, 43.
Gastrula, 43.
-stage in star-fish, 60.
Giant cells, action on tubercle bacilli,
162.
origin of, 135, 159.
in Arthropoda, 79.
Giard, 82.
INDEX.
215
' Glande indeterminee,' 78.
Gley, 153.
Goloubew, 138.
Gonococci, behaviour of leucocytes to,
117.
Gordtus, parasitic on Nais, 67.
Granulation-tissue, origin of, 132.
Granuloma, formation of, 1 58.
Grawitz, 9, 132.
Gregarina, parasite of Earth-worm, 68.
Griesbach, on phagocytosis in Molluscs,
77-
Grobben, 66.
Gryllidse, 88.
,, Gastrsea theory, 43.
on ingestion of granules by leuco-
cytes, 76.
Hafkine, epidemic in Paramsecia, 21.
Hankin, 176.
Hanstein, on lesions of alga;, 16.
Hemitis, 186.
Hepatic capillaries, endothelium of, 139.
Hering, mechanical theory of inflam-
mation, 144.
Hermann, 81.
Hertwig, 156.
Hess, 126.
Hircinia echinata, 52.
Hofer, on digestion in Amcebse, 19.
Hofmeister, peptone in leucocytes, 124.
Hydra, reaction to injuries, 57.
Hydromedusse, 58.
Hydrotropism in plasmodia, 39.
Hymenoptera, 88.
Hypersemia, 5.
Hypertrophic cirrhosis, 170.
IBENER, 176.
Immunity, 156.
Infectious diseases, absence in Sponges,
54-
of Infusoria, 2, 17 28.
Inflammation, author's definition of,
189.
Cornil and Ranvier's definition of,
8.
Neumann's definition of, 1 1 .
Ziegler's definition of, 7.
theories of, 5, 180.
biological theory of, 184.
paralytic theory of, 6.
spasmodic theory of, 6.
Virchow's theory of, 6, 1 80.
Samuel and Cohnheim's views, 6,
181.
evolution of, 109.
in plants, 39.
in Triton embryo, 98 103.
Inflammation without diapedesis, 186.
Inflammations, chronic, 157 171.
serous, 171 179.
Infusoria, artificial lesions in, 15 17.
biological relations between, 2.
colonies of, 44.
infectious diseases of, 17 28.
Insects, varieties of phagocytes in, 88.
Intracellular digestion in Molluscs, 78.
Invertebrata, leucocytes of, 76.
Isaria destructor, 90.
Ischikawa, on Hydra, 57.
'ftio in paries? 147.
JEQUIRITY, action on leucocytes, 119.
KARYOKINESIS, 9, 10.
in connective tissue cells, 97.
Kebner, 169.
Keratitis, experimental, 8.
Kitasato, 177.
Klebs, 138.
on infection of Euglena, 24.
Klemperer, on antitoxic properties of
serum, 177.
Koch, 125.
giant cells, 167.
Korschelt, 61.
Koultchitsky, 130.
Kowalewski, on phagocytes of Asci-
dians, 79.
on spleen of Molluscs, 77.
Krukenberg, peptic ferment in Myxo-
mycetes, 30.
on a tryptic ferment in Sponges,
5-
Kupffer, on star-cells, 140.
LANDEREK, 150.
Lange, 117.
Leber, chemiotaxis of leucocytes, 117.
digestive ferment in pus, 124.
Le Dantec, digestion in Stentor, 20.
Lendenfeld, de, 55.
physiology of Sponges, 49.
Lepidoptera, 88.
Leprosy, 168.
Leprosy bacilli, behaviour of leucocytes
to, 117.
ingestion by endothelial cells, 140.
Leptotrix, digestion of, 19.
in young Spongillae, 52.
Lesions, traumatic, in lower animals,
14.
Lethargy of silkworms, 89.
Leucin, action on leucocytes, 119.
Leuchsemia, 114.
Leucocytes, in 136.
2l6
INDEX.
Leucocytes, emigration of, 144.
in tubercle, 159.
ofinvertebiata, 76.
origin of pus-cells from, 9.
peripheral distribution of, 147.
psychical activity of, 192.
varieties of, in.
Leucocytosis after injection of pro-
teins, 121.
Ley dig, 138.
Lieberkiihn, 114.
digestion of Infusoria in Sponges,
47-
Limbeck, von, on Ieucocytosis,i2i.
Lubarsch, 118, 121, 176.
on reactive phenomena in Asci-
dians, 80.
Luminous bacteria as parasites, 82.
Lupus, 1 66.
Lymphocytes, 114.
in tubercle, 159.
MACROPHAGES, 161.
as scavengers, 190.
Malaria, endothelial cells in, 141.
Marchand, 132.
Marmot, tubercle of, 1 60.
Massart and Bordet, irritability of
leucocytes, 118, 145.
' Mastzellen,' 143.
Medusas, inflammation in, 59.
Meriones, tubercle in, 162.
Merotomy of Amoeba, 14.
Mesoderm, formation of, in star-fish, 61.
Mesostomum Ehrenbergl, 66.
Microbicidal action of serum, 174.
Microsphsera, 17.
Microsporidium of ' pebrine,' 92.
Migratory cells, conversion into con-
nective tissue cells, 106.
Mollusca, 75 93.
Monads, digestion in flagellated, 19.
infection of, 25.
Monocystis (Gregarinse) 68.
Monospora bicuspidata, 83.
Mucor helminthophorus , 73.
Miiller, epidemic in Paramsecia, 20.
Muscardine, 90.
Muskatbluth, 130, 169.
Myelocytes, 114.
Myxomycetes, plasmodium of, 29.
Nais proboscidea ,67.
Neisser, 140, 169.
Nematoda, 70 74.
producing tubercles, 170.
Nerves, their influence in inflamma-
tion, 150.
Netchaeff, on litmus-granules in leu-
cocytes, 124.
Neumann, on inflammation, u.
Neutrophile leucocytes, 113.
Nikiforoff, 131.
Nissen, 175.
Nucleus, importance for regeneration,
IS-
(EDEMA, inflammatory, 173.
Olpidium, 25.
Orthoptera, 88.
Oryctes, 78.
Ostracoda, 75.
Ouskoff, 114.
Pandorina morum, 25 .
Paralytic theory of inflammation, 6.
Paramaecia, epidemic in, 20.
Parasitism in Paramaecia, 20.
of Infusoria, 2.
Parenchymatous inflammation, 9.
Pasteur, on diseases of silkworms, 89.
Pasteuria ramosa, 87.
Pathology, necessity for comparative, 4,
12, 195-
1 Ptbrine ' infecting Daphnia, 87.
of silkworms, 92.
Pee, on leucocytosis, 1 21.
Pekelharing, 1 1 8.
Peremeschko, 130.
Pericardial phagocytes of insects, 88.
Peritoneal endothelium, phagocytic
nature of, 66.
Perivisceral cells in Worms, 66.
Petruschky, 125.
Peziza sclerotiorum, 41.
Pfeffer, on chemiotaxis, 33.
Phagocytella, 43.
Phagocytes in Hydromedusse, 58.
their absence in certain Crustacea,
87.
fate of, 107.
in insects, 88.
in tubercle, 159.
Phagocytic properties of leucocytes,
114.
Phagocytosis in Annelida, 66, et seq.
in Crustacea, 78.
in Daphnia, 84.
in recurrent fever, 1 86.
in Sponges, 47, 51.
in star-fish, 61.
Philina, 77.
Philliroe, 77.
Physarum, 31, 36.
Piorry, 186.
Plants, inflammation in, 39.
INDEX.
217
Plasma-cells, 143.
Plasmodia, formation of, in Bipinnaria,
63-
formation of, in Annelida, 67.
formation by fusion of cells, 52.
of Myxomycetes, 29.
Pleurobranchaea, 77.
Pneumonia, 177.
Polyphagus Euglencc, 25.
Ponfick, 140.
Protospongia, 45.
Protozoa, digestion in, 18.
reactive phenomena of, 15 28.
Pseudo-eosinophile leucocytes, 113.
Pseudo-navicellae, 69.
Pus-cells, origin of, 9.
QUININE, action on leucocytes, 119,
145.
RANVIER, on inflammation, 8.
division of leucocytes, 129.
endothelial cells of omentum,
142.
Rat's serum, bactericidal action of, 176.
Reaction of digesting cells of Spongilla.
So-
Recklinghausen, emigration of leuco-
cytes, 149.
Recurrent fever, 186.
Reparation in inflammation, 10.
Rhabditis invading Earthworm, 70.
Rhizopoda, digestion in, 19.
Jfhisostomum Cuvieri, 59.
Ribbert, 132.
lymphatic glands, 142.
Roeder, 176.
Roemer, 119.
Roger, 151.
Roser, views on inflammation, 10,
no.
Rossbach, 123.
Roux, 176.
and Yersin, on pleurisy in diph-
theria, 173.
SACHS, on inflammation, u.
Samuel, 147, 150.
theory of inflammation, 6, 181.
Sandhoppers, fungoid disease of, 8l.
Saprolegnise infecting Crustacea, 82,
86.
Saville Kent, inclusion of bacteria by
Physarum, 38.
on Protospongia, 45.
Sawtchenko, 117, 169.
Scavenger function of leucocytes, 190.
Schafer, 127.
Schueppel, 162.
Schultze, Max, 113.
Selenka, 61.
Sensibility of leucocytes, 117.
tactile, of leucocytes, 123.
Serous exsudation, functions of, 174,
177.
Serous inflammations, 171 179.
Serum, bactericidal action of, 175.
Severini, 138.
Silkworms, diseases of, 89, 91.
Siphonochalina coriacea, 47.
Snellen, 152.
Soudakewitch, 166, 169.
Spasmodic theory of inflammation, 6.
Spermatozoids of ferns, chemiotaxis in,
34-
Spermophilus, tubercle of, 160.
Sphserophrya magna, 3.
Sphxrophrya paramsecioruin, 3.
Spiriila, 186.
Spirobacillus Cienkowskii, 87.
Spleen of Gasteropoda, 77.
Sponges, 4555-
effects of wounding, 51.
phagocytosis in, 47, 51.
Spongilla, digestion in, 48.
Spores, fate of, in leucocytes, 128.
Sporozoa infecting Crustacea, 82, 87.
infecting insects, 92.
Spronck, on leucocytes, 130.
Spumaria alba, 148.
Stahl, on plasmodia of Myxomycetes,
33-
Star-cells, 140.
' Staubzellen,' 160.
Stentor, merotomy of, 16.
Stephanos typ/ms, 53.
Stern, 174.
Stigmata, 138.
Stomata, 138.
Strieker, no, 138.
Stylonychia, 22.
Swine septicaemia in pigeons, 141.
TADPOLES, inflammation in, 104.
Talitrus (sandhopper), 81.
Tchistowitch, 135, 160.
Temperature in inflammation, 12.
Ttrebella, 66.
Tetanus, 177.
Theories of inflammation, 5, 180.
Thethys, 76.
Thiocystis, 20.
Thoma, on inflammation, II.
Tissue-cells, origin of pus-cells from, 9.
Tizzoni, 177.
Teuton, 140.
Trachelius ovum, 16.
Trapeznikoff, 129.
218
INDEX.
Traumatic lesions in lower animals,
14.
Triton tseniatus, 98.
Trophotropism, 33.
Tubercle bacilli, degeneration of, 161.
Tuberculin, injection of, 168.
effect on guinea-pigs, 178.
Tuberculosis, 157 168.
avian, 161.
zooparasitic, 170.
Tunicata, 75 93.
Turbellaria, phagocytic reaction in,
66.
URODELA, inflammation in, 95.
Uromyces Pisi, 41.
VACCINATION, effect on inflammatory
reaction, 154.
Vacuolar degeneration, 169.
Vaillard, 177.
Vascular system in inflammation, 8.
in invertebrata, 75.
Vasomotor nerves in inflammation,
I5I-
Vaucheria, recovery from injury, 17.
Vermes, 65 74.
Vertebrata, inflammation in, 94 197.
Vesuvine, reaction of ingested bacilli
to, 19.
Vibrio Metchnikywii, 154, 178.
Virchow, theory of inflammation, 6,
1 80.
WALDENBURG, inflammation in
plants, 39.
Weber's law applied to lower animals,
34-
Weigert, giant cells, 166.
origin of pus-cells, 9.
Worms, inflammation in, 65 74.
Wyssokowitch, 140.
YERSIN, 135.
ZIEGLER, on inflammation, 7.
origin of connective tissue, 131.
Zimmerman, 173.
Zoochlorellx, 53.
Zooparasitic tuberculosis, 170.
Zoox i ntellse, 53.
Zopf, 73.
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